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Accepted by O. Seeman: 19 Mar. 2015; published: 13 Apr. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
Zootaxa 3946 (4): 567
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576
www.mapress.com
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zootaxa
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Article
567
http://dx.doi.org/10.11646/zootaxa.3946.4.5
http://zoobank.org/urn:lsid:zoobank.org:pub:0252EEB7-3576-456E-9E55-240221B7258E
Two new feather mites of the genus Neocalcealges Orwig
(Analgoidea: Trouessartiidae) from the Sichuan province of China
ZI-YING WANG
1, 2
& HEATHER PROCTOR
2, 3
1
Institute of Entomology, College of Plant Protection, Southwest University, Chongqing, China 400715. E-mail: zhibaoy@swu.edu.cn
2
Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada T6G 2E9. E-mail: hproctor@ualberta.ca
3
Corresponding author
Abstract
Two new species of the feather mite genus Neocalcealges Orwig 1968 (Analgoidea: Trouessartiidae) are described from
passerine birds (Passeriformes) from the Sichuan province of China: Neocalcealges davidi sp. nov. from Alcippe davidi
Styan (Leiothrichidae) and N. chrysotis sp. nov. from Lioparus chrysotis (Blyth) (Paradoxornithidae). We provide a key
to all described species of Neocalcealges.
Key words: bird mites, Acari, Astigmata, new taxa
Introduction
The feather mite family Trouessartiidae includes 11 genera, most of which are associated with perching birds
(Passeriformes) (Gaud & Atyeo 1996). The genus Neocalcealges Orwig, 1968 was established in the revision of
selected genera of Trouessartiidae and originally contained six species (Orwig 1968). Together with four other
trouessartiid genera it is characterized by having the ambulacra of legs III and IV asymmetrically developed.
Neocalcealges is morphologically most similar to Calcealges Gaud, 1952 but differs in the following characters:
setae d2 are setiform and short in Neocalcealges vs long and strong in Calcealges; coxal apodemes I are fused into
a "V" or "U" shape in both sexes vs free in Calcealges; and coxal fields III are open in males vs closed in
Calcealges (Gaud 1952; Orwig 1968).
Of the six species that have been assigned to date to the genus Neocalcealges, five were collected from
laughing thrushes and babblers (Leiothrichidae and Timaliidae), and N. cuspilobus Orwig 1968 was collected from
fairy flycatchers (Stenostiridae) as well as from hosts of the two above-mentioned families (Orwig 1968). Herein,
we add two new species of Neocalcealges from passerines of the families Leiothrichidae and Paradoxornithidae in
China and provide a key to all described species.
Material and methods
Birds were captured in Dujiangyan City, Sichuan Province, China by ornithologists working under corresponding
permits issued by Chinese authorities. One of us (ZW) removed mites from two bird specimens, which accidentally
died during the process of capturing, and placed them in tubes with 96% ethanol. Mites were cleared in lactic acid,
slide-mounted in Hoyer’s medium, and dried for 4 days at 50°C on a slide warmer. Drawings were made at 200× or
400× using a 1× a drawing tube attached to a Leica DMLB compound microscope with differential interference
contrast (DIC) lighting. Measurements were made at 200× and 400× using a calibrated ocular micrometer.
In species descriptions, all measurements are given in micrometers (μm). Idiosomal length was measured from
the anterior margin of the propodonotum to the posterior end of the opisthosomal lobes. Widths of the idiosoma
and hysteronotal shield were measured at the level of setae cp. The length of the propodonotal shield was measured
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along the median line of the shield, and the width was measured at its widest part. The length of the hysteronotal
shield was measured along its lateral border, and the width was measured at its widest part, at the level of setae cp.
The terminology relating to the idiosomal setation follows Griffiths et al. (1990) with modifications of coxal
setae 3a to 4b, and 3b to 3a following Norton (1998). The leg setation follows Grandjean (1939). The descriptions
follow the standard format used for mite species of the family Trouessartiidae (Orwig 1968; Mironov & González-
Acuña 2013). Holotypes (male) and at least one male and female paratype for both species described here are
deposited in the Institute of Zoology, Chinese Academy of Sciences, Beijing, China (IOZ). Paratypes are also
deposited at the Institute of Entomology, Southwest University, Chongqing, China (IESWU); Zoological Institute
of the Russian Academy of Sciences (Saint-Petersburg, Russia) (ZISP) and the Strickland Entomological Museum
at the University of Alberta (Edmonton, Canada) (UASM). Host systematics follows Clements et al. (2014).
TABLE 1. Hosts and locations for Neocalcealges species. References: 1 = Orwig (1968), 2 = McClure et al. (1973).
Mite species Host species or subspecies in Clements et al. (2014) and original mite-
host record reference
Location collected
N. angustus Cyanoderma erythropterum (Blyth) (Timaliidae) [=Stachyris
erythroptera in 1,2]
Gombak, Malaya
Malacopteron cinereum (Eyton) (Pellorneidae) [1,2] Malaya
N. chrysotis
sp. nov. Lioparus chrysotis (Blyth) (Paradoxornithidae) Sichuan, China
N. cuspilobus Alcippe nipalensis (Hodgson) (Leiothrichidae) [2] Malaya
Alcippe peracensis peracensis Sharpe (Leiothrichidae) [1] Malaya
Culicicapa ceylonensis (Swainson) (Stenostiridae) [1,2] Gombak, Malaya
Stachyris nigriceps Blyth (Timaliidae) [1,2] Malaya
N. davidi
sp. nov. Alcippe davidi Styan (Leiothrichidae) Sichuan, China
N. emarginatus Ianthocincla mitrata (Müller) (Leiothrichidae) [=Garrulax mitratus in
1,2]
Pahang and Selangor,
Malaya
Leiothrix argentauris (Hodgson) (Leiothrichidae) [2] Malaya
N. inauditus Actinodura cyanouroptera sordidior (Sharpe) (Leiothrichidae) [=Minla
cyanouroptera sordidior in 1]
Pahang, Malaya
Actinodura cyanouroptera (Hodgson) (Leiothrichidae) [=Minla
cyanouroptera in 2]
Malaya
Actinodura strigula (Hodgson) (Leiothrichidae) [=Minla strigula in 1,2] Thailand; Malaya
N. segregatus Actinodura cyanouroptera (Hodgson) (Leiothrichidae) [=Minla
cyanouroptera in 2]
Malaya
Actinodura cyanouroptera sordidior (Sharpe) (Leiothrichidae) [=Minla
cyanouroptera sordidior in 1]
Malaya
Heterophasia picaoides (Hodgson) (Leiothrichidae) [2] Malaya
Ianthocincla mitrata (Müller) (Leiothrichidae) [=Garrulax mitratus in
1,2]
Malaya
Leiothrix argentauris (Hodgson) (Leiothrichidae) [1,2] Malaya
Rhipidura albicollis (Vieillot) (Rhipiduridae) [2] Malaya
Stachyris nigriceps davisoni Sharpe (Timaliidae) [1] Pahang and Genting
Sempah Malaya;
Pattalung, Thailand
Stachyris nigriceps Blyth (Timaliidae) [2] Malaya
Stachyris poliocephala (Temminck) (Timaliidae) [1,2] Malaya
N. undulatus Heterophasia picaoides (Hodgson) (Leiothrichidae) [1,2] Pahang, Malaya
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TWO NEW SPECIES OF FEATHER MI TES OF NEOCALCEALGES
Systematics
Family Trouessartiidae Gaud, 1957
Genus Neocalcealges Orwig, 1968
Neocalcealges davidi sp. nov.
(Figs. 1, 2, 3A, 4A–D)
Description. MALE (holotype). Body 430 long (400–430 in 6 paratypes) and 210 wide (190–220). Dorsum.
Distance between prodorsal and hysteronotal shields 38 (20–33). Prodorsal shield: 130 long (120–130) and 133
wide (130–140), surface without lacunae, but each posterolateral corner of shield with two ovate lightly sclerotized
patches; lateral areas not encompassing bases of setae se, not fused with scapular shields. Setae si short setiform,
12 (16–20) in length, separated by 49 (50–53); setae se long, about 160 long. Humeral shield with setae c2
lanceolate, 41 (36–41) in length. Hysteronotal shield length 240 (230–240), width 140 (130–140), surface with
some almost transparent lightly sclerotized patches, number and shape of patches not consistent among individuals.
Setae c1 absent. Setae d1 and e2 present. Lateral margin of hysteronotal shield with pair of wide and shallow
incisions at level of trochanters IV and pair of small and deep lateral incisions at level of setae e2 extending to
bases of these setae. A pair of narrow curved platelets present on lateral parts of hysteronotum at level of
trochanters III, possibly analogous to metapodosomal shields of Park and Atyeo (1971). Setae f2 present only in
form of indistinct alveoli near base of setae ps2, apparently placed laterally rather than dorsally. Setae h1 situated
anterior to setae h2. Terminal lamellae with 6 (4–7) small irregular indentations, creating 7 (5–8) teeth. Ve n t e r .
Coxal apodemes I fused into a U. Setae c3 short lanceolate, 26 (23–30) in length. Coxal fields of legs III open.
Coxal apodeme IIIa+ IV long and continuous. Genital apparatus situated at middle level of coxal apodemes IVa.
Adanal apodemes well developed, but apices not contacting coxal apodeme IVa. Distance between setae: g:ps3 12
(11–14); g:g 24 (19–26). Translobar apodeme present. A pair of cupules ih located near posterior ends of
opisthosomal lobes. Legs. Ambulacral discs of legs III and IV wider than those of legs I and II and asymmetrical.
Setae sRIII setiform, about 12 in length. Setae d and e of tarsi IV barrel-shaped, with discoid cap, situated
subapically.
FEMALE (7 paratypes). Body 460–510 long and 200–230 wide. Dorsum. Distance between prodorsal and
hysteronotal shields about 21–32. Prodorsal shield: 120–140 long and 140–160 wide, surface same as in males,
each posterolateral corner of shield with two ovate lightly sclerotized patches; lateral areas not encompassing bases
of setae se, not fused with scapular shields. Setae si short setiform, 21–27 in length, separated by 51–57; setae se
long, about 170 long. Scapular shields very narrow. Humeral shield with setae c2 lanceolate, 34–38 in length.
Hysteronotal shield: length 270–300, width 140–160, surface with some almost transparent lightly sclerotized
patches, number and shape of patches not consistent among individuals. Setae c1 missing. Setae d1 and e2 present.
Lateral margins of hysteronotal shield sinuous. Setae f2 situated on membranous integument adjacent to setae h2.
Setae h1 situated anterior to setae h2 and posterior to supranal concavity. Setae ps1 located on dorsal surface,
anterior to setae h3. Copulatory aperture dorsal, anterior to setae e2, 42–57 from supranal concavity. Distance from
h3 to tips of opisthosomal lobes 39–47. Ven te r. Coxal apodemes I fused into a U. Setae c3 short lanceolate, 24–32
in length. Coxal fields of legs III open. Epigynum well-developed, enveloping setae 4b, g and genital discs.
Distance between setae: 4b:g, 30–37; g:g, 86–98; 4b:4b, 33–42. Coxal apodemes IVa present as small sclerites.
Legs. Ambulacral discs of legs III and IV wider than those of legs I and II and asymmetrical. Setae sRIII setiform,
about 35 in length.
Type material. Holotype male [IOZ(E) 227734], 6 male and 7 female paratypes ex David’s Fulvetta, Alcippe
davidi Styan (Passeriformes: Leiothrichidae), CHINA: Sichuan, Dujiangyan, 31°14′07''N, 103°35′16''E, 20 May
2014, coll. H.-Y. Cao.
Etymology. The specific epithet derives from the specific name of the host and is a noun in apposition.
Differential diagnosis. Among known species of the genus Neocalcealges, the new species is closest to N.
emarginatus Orwig, 1968 from the Chestnut-capped Laughingthrush Ianthocincla mitrata (Müller) (Passeriformes:
Leiothrichidae) (Orwig 1968). In both sexes of the two species, setae d1and e2 are present. In males of both
species, one pair of narrow curved platelets (~metapodonotal platelets) are present at the level of trochanters III,
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branching from the lateral margins of the hysteronotal shield, and the terminal lamellae have indentations. The new
species differs from N. emarginatus in the following characters: in males of N. davidi sp. nov., the lateral margin of
hysteronotal shield has a pair of shallow incisions at the level of coxae IV and a pair of narrow and deep lateral
incisions at the level of setae e2 extending to bases of these setae; the platelets at the level of trochanters III are
long and extend to the lateral margins of the hysterosoma; setae f2 are reduced to alveoli near the base of setae ps2;
the terminal lamellae have six (4–7) indentations; in females, setae h1 are posterior to the supranal concavity; the
opisthosomal lobes are long, 39–47 in length. In males of N. emarginatus, the lateral incisions of the hysteronotal
shield at the level of setae e2 are shallow and not extending to the bases of these setae; the platelets at the level of
trochanters III do not extend to the lateral margin of the body; setae f2 are present; the terminal lamellae have only
two indentations; in females, setae h1 are anterior to the supranal concavity; the opisthosomal lobes are short, about
25 in length. Orwig (1968) states that males of Neocalcealges have one pair of small pores, cupules ip, located on
the margins of opisthosoma at the level of translobar apodeme and a second pair, cupules ih, near the posterior ends
of the opisthosomal lobes; however, we were unable to see cupules ip on the margins of opisthosoma in our
specimens.
FIGURE 1. Neocalcealges davidi sp. nov., male. A—dorsal view, B—ventral view.
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TWO NEW SPECIES OF FEATHER MI TES OF NEOCALCEALGES
FIGURE 2. Neocalcealges davidi sp. nov., female. A—dorsal view, B—ventral view, C—base of spermatheca; ca –
copulatory aperture.
Neocalcealges chrysotis sp. nov.
(Figs. 3B, 4E, 5–6)
Description. MALE (holotype). Body 435 long (410–440 in 8 paratypes) and 200 wide (200–210). Dorsum.
Distance between prodorsal and hysteronotal shields 8 (3–18). Prodorsal shield: 140 long (140–150) and 150 wide
(150–160), surface uniformly punctate without lacunae or poorly sclerotized patches; lateral areas not
encompassing bases of setae se, not fused with scapular shields. Setae si short setiform, 19 (23–27) in length,
separated by 59 (55–65); setae se long, 170 (150–170) long. Humeral shield with setae c2 lanceolate 49 (49–57) in
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length. Hysteronotal shield length 270 (240–270), width 150 (150–160). Surface without lacunae. Setae c1
missing. Setae d1 and e2 present. Lateral margin with pair of wide and shallow incisions at level of trochanters IV
and pair of narrow and deep lateral incisions at level of setae e2 extending to bases of these setae. A pair of narrow
and bent platelets present at level of trochanters III, not connected to hysteronotal shield, possibly analogous to
metapodosomal shields of Park and Atyeo (1971). Setae f2 represented only as very faint alveoli near base of setae
ps2 in some specimens or not visible at all. Setae h1 situated anterior to setae h2. Terminal lamellae each with 3
indentations. Vent e r. Coxal apodemes I fused into a V. Setae c3 short lanceolate, 25 (22–30) in length. Coxal fields
of legs III open. Coxal apodemes IIIa+ IV long, short anterior parts of apodemes IIIa+ IV split from remaining
parts of these apodemes. Genital apparatus situated at middle level of coxal apodemes IVa. Adanal apodemes well
developed, apices touching coxal apodemes IVa. Distance between setae: g:ps3 10 (9–10); g:g 18 (19–24).
Translobar apodeme present. Cupules ih located near posterior ends of opisthosomal lobes. Legs. Ambulacral discs
of legs III and IV wider than those of legs I and II and asymmetrical. Setae sRIII setiform, 53 (41–58) in length.
Genua IV with paraxial, semicircular membranes. Setae d and e of tarsi IV barrel-shaped, with discoid cap, situated
subapically.
FIGURE 3. Neocalcealges davidi sp. nov., male. A—ventral view of opisthosoma; Neocalcealges chrysotis sp. nov., male.
B—ventral view of opisthosoma.
FEMALE (6 paratypes). Body 480–530 long and 190–220 wide. Dorsum. Distance between prodorsal and
hysteronotal shields 7–22. Prodorsal shield: 140–150 long and 160–170 wide, surface same as in males,
posterolateral surface without lacunae or poorly sclerotized patches; lateral areas not encompassing bases of setae
se, not fused with scapular shields. Setae si short setiform, 22–35 in length, separated by 60–70; setae se long, 150–
170 long. Scapular shields very narrow. Humeral shield with setae c2 lanceolate 47–56 in length. Hysteronotal
shield length 300–320, width 150–160, surface with a few lightly sclerotized patches near anterior margin. Setae c1
missing. Setae d1 and e2 present. Lateral margin of hysteronotal shield gently undulating. Setae f2 situated on
membranous integument adjacent to setae h2. Setae h1 situated anterior to setae h2. Setae ps1 located on dorsal
surface, anterior to setae h3. Copulatory aperture dorsal, anterior to setae e2, 62–75 from supranal concavity.
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Distance from h3 to tips of opisthosomal lobes 39–41. Ve n t e r. Coxal apodemes I fused into a V. Setae c3 short
lanceolate, 24–30 in length. Coxal fields of legs III open. Epigynum well developed, enveloping setae 4b, g and
genital discs. Distance between setae: 4b:g 38–41; g:g 93–110; 4b:4b 30–45. Coxal apodeme IVa absent. Legs.
Ambulacral discs of legs III and IV wider than those of legs I and II and asymmetrical. Setae sRIII setiform, 31–39
in length.
FIGURE 4. Neocalcealges davidi sp. nov., legs of male. A- D— legs I-IV in dorsal view; Neocalcealges chrysotis sp. nov.,
E—leg IV of male in dorsal view; mem— membrane.
Type material. Holotype male [IOZ(E) 227735], 8 male and 6 female paratypes ex the Golden-breasted
Fulvetta Lioparus chrysotis (Blyth) (Passeriformes: Paradoxornithidae), CHINA: Sichuan, Dujiangyan,
31°14′21''N, 103°35′14''E, 16 May 2014, coll. H.-Y. Cao.
Etymology. The specific epithet derives from the specific name of the host and is a noun in apposition.
Differential diagnosis. The new species N. chrysotis sp. nov. is closest to N. davidi sp. nov. from Alcippe
davidi Styan (Leiothrichidae) described above. In both sexes of the two species, setae d1and e2 are present. In
males of both species, the shapes of hysteronotal shield are similar, the lateral margins have a pair of shallow
incisions at the level of coxae IV and a pair of narrow and deep lateral incisions at the level of setae e2; setae f2 are
absent or reduced to faint alveoli, and the terminal lamellae have indentations. The new species differs from N.
davidi in the following characters: in both sexes of N. chrysotis, the distance between the prodorsal and
hysteronotal shields is short (usually less than 20 µm); coxal apodemes I are fused into a V; in males of N.
chrysotis, a pair of narrow platelets (~metapodonotal platelets) are present at the level of trochanters III but not
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connected to the hysteronotal shield; the terminal lamellae have 3 indentations; small anterior parts of coxal
apodemes IIIa+ IV are split from remaining body of these apodemes; genua IV have paraxial, semicircular
membranes; in females, the distance from the copulatory aperture to the supranal concavity is long, 62–75 µm. In
both sexes of N. davidi, the distance between the prodorsal and hysteronotal shields is long (usually over 20 µm);
coxal apodemes I are fused into a U shape; in males of N. davidi, the narrow platelets situated at the level of
trochanters III (~metapodonotal platelets) are connected to the hysteronotal shield; the terminal lamellae have 4–7
indentations; coxal apodemes IIIa+ IV are long and continuous; in females, the distance from the copulatory
aperture to the supranal cleft is short, 42–57 µm.
FIGURE 5. Neocalcealges chrysotis sp. nov., male. A—dorsal view, B—ventral view.
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FIGURE 6. Neocalcealges chrysotis sp. nov., female. A—dorsal view, B—ventral view, C—base of spermatheca.
Key to species of Neocalcealges (modified from Orwig 1968)
1. Setae e2 absent; setae d1 present or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Setae e2 and d1 present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Setae d1 absent. Male: distance between setae g and ps3 7–12 µm. Female: copulatory aperture located far from anterior end of
supranal cleft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cuspilobus Orwig, 1968
- Setae d1 present. Male: distance between setae g and ps3 13–17 µm. Female: copulatory aperture located close to anterior end
of supranal cleft. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. undulatus Orwig, 1968
3. Male: posterior margin of terminal lamellae margins smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Male: posterior margin of terminal lamellae margins with indentations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4. Male: distance between setae g and ps3 20 µm or more. Female: setae h1 located anterior to end of supranal concavity, far from
setae h2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
- Male: distance between setae g and ps3 10 µm or less. Female: setae h1 located posterior to anterior end of supranal concavity,
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closed to setae h2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. inauditus Orwig, 1968
5. Male: distance between setae g 23–29 µm, terminal lamellae rounded. Female: distance between supranal cleft and copulatory
aperture 12–18 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. segregatus Orwig, 1968
- Male: distance between setae g 15–17 µm, terminal lamellae truncate. Female: distance between supranal cleft and copulatory
aperture 29–33 µm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. angustus Orwig, 1968
6. Male: terminal lamellae each with 3 or more indentations. Female with setae h1 located posterior to end of supranal cleft, close
to setae h2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- Male: terminal lamellae each with 2 indentations. Female with setae h1 located anterior to end of supranal cleft, far from setae
h2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. emarginatus Orwig, 1968
7. Male: terminal lamellae each with 4–7 indentations (Fig. 3A). Female:distance from copulatory aperture to supranal concavity
42–57 µm (Fig. 6A). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. davidi sp. nov.
- Male: terminal lamellae each with 3 indentations (Fig. 3B). Female: distance from copulatory aperture to supranal concavity
62–75 µm (Fig. 2A). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. chrysotis sp. nov.
Acknowledgements
The research was supported by the National Natural Science Foundation (31401994); the Fundamental Research
Funds for the Central Universities (XDJK2014C182) and the State Scholarship Fund of China Scholarship Council
(201306995039) (all to ZW) and a Natural Sciences and Engineering Research Council of Canada Discovery Grant
to HP .
References
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Clements checklist of birds of the world: Version 6.9. Available from: http://www.birds.cornell.edu/clementschecklist/
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Gaud, J. (1952) Sarcoptides plumicoles des oiseaux de Madagascar. Mémoires de l'Institut Scientifique de Madagascar, Série A,
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