Corresponding author: Michael Staab (
Academic editor: M. Borowiec
The genus
Staab M, Hita Garcia F, Liu C, Xu Z-H, Economo EP (2018) Systematics of the ant genus
Recent phylogenetic studies have clarified the evolutionary history of ant subfamilies and genera. One higher-level taxon consistently recovered is the subfamily
The genus has been comprehensively revised on a global scale by
In the last decade, X-ray microtomography (
Despite its common usage in invertebrate paleontology, as well as functional and comparative morphology (e.g.
In this study, we provide a review of the genus
The collection abbreviations follow
The material of the new species was collected during recent ecological field work activities of the first author (see e.g.
The following measurements (all expressed in mm) and indices are based on
We scanned all Chinese
Overview of
Species | Identifier | Type status | Magnification (x) | Exposure (s) | Voxel size (µm) | Source distance (mm) | Detector distance (mm) | Voltage (kV) | Power (W) | Amperage (µA) |
---|---|---|---|---|---|---|---|---|---|---|
|
|
non-type | 4 | 1.5 | 4.826 | 24.99 | 10 | 45.24 | 3.54 | 78.14 |
|
|
holotype | 4 | 1.5 | 4.359 | 19.99 | 11 | 45.23 | 3.54 | 78.3 |
|
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holotype | 4 | 1.5 | 4.359 | 19.99 | 11 | 45.24 | 3.55 | 78.35 |
|
OKENT0016142 | non-type | 4 | 0.8 | 3.660 | 13 | 11 | 45.24 | 3.54 | 78.29 |
|
|
non-type | 4 | 0.8 | 3.897 | 14.99 | 11 | 45.24 | 3.54 | 78.17 |
|
|
non-type | 4 | 0.8 | 3.097 | 11 | 13 | 45.24 | 3.54 | 78.19 |
|
|
paratype | 4 | 1.5 | 2.534 | 12 | 19.99 | 45.24 | 3.54 | 78.19 |
|
|
holotype | 4 | 1.5 | 4.193 | 17.99 | 11 | 45.24 | 3.53 | 78.05 |
|
|
paratype | 4 | 1.5 | 2.252 | 11 | 22 | 45.24 | 3.54 | 78.33 |
All specimens used in this study have been databased and the data are freely accessible on AntWeb (
=
=
This key is partly derived from
1 | In profile, petiolar node squamiform and rectangular, high and erect (Fig. |
|
– | In profile, petiolar node never squamiform, either low, elongate, and barrel-shaped, or rounded-triangular (Fig. |
|
2 | In profile, petiolar node clearly narrowing dorsally, broader on the base than on the apex (Fig. |
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– | In profile, petiolar node not or only weakly narrowing dorsally, the base as or almost as broad as the apex (Fig. |
|
3 | Anterior clypeal margin with a distinct and broad notch (Fig. |
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– | Anterior clypeal margin without a distinct and broad notch (Fig. |
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4 | Frontal carinae weakly developed, short, little diverging above antennal insertions, and with narrow lateral lamellae; dorsal surface of body without erect hairs protruding from dense pubescence (Fig. |
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– | Frontal carinae better developed, long, diverging above antennal insertions, and usually with broad lateral lamellae; dorsal surface of body with erect hairs protruding from dense pubescence (Fig. |
|
5 | In profile, posterodorsal corners of propodeum rounded (Fig. |
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– | In profile, posterodorsal corners of propodeum angular (Fig. |
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6 | Scapes without erect hairs protruding from the dense pubescence; frontal carinae touching each other at their anteriormost level (Fig. |
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– | Scapes with many erect hairs protruding from the dense pubescence; frontal carinae clearly separated at their anteriormost level, not touching each other, their lamellae broad, conspicuously extending laterally above antennal insertions (Fig. |
|
7 | Propodeal declivity punctured, mostly opaque; frontal furrow conspicuous and darker than anterior cephalic dorsum (Fig. |
|
– | Propodeal declivity very shiny, at most superficially punctured; frontal furrow inconspicuous and of same color than anterior cephalic dorsum (Fig. |
|
Petiole in profile view.
Petiole, abdominal segment III and anterior portion of abdominal segment IV in profile view.
Anterior portion of cephalic dorsum, in full-face view (
Mesosoma dorsum in profile view.
Propodeum and petiole in profile.
Anterior portion of cephalic dorsum in full-face view.
Anterior portion of cephalic dorsum in full-face view (
Workers of this clade can be separated from all other
This clade includes seven species and is restricted to east and southeast Asia. All species except
Still images from surface display volume renderings of 3D model of
In full-face view, head slightly longer than broad (
Mesosoma in profile slightly convex and as long as maximum head length including mandibles (
Petiolar node in profile high, nodiform, with a straight and sloping anterior face, dorsum of node broadly rounded, posterior face as steep as anterior face and relatively short, less than half as long as anterior face; petiole in dorsal view longer than broad, apex of node almost as long as broad; ventral process of petiole well developed, with a roughly trapezoid projection of varying shape and ventral outline (see ‘variation’).
In dorsal view abdominal segment III anteriorly much broader than petiole; its sides convex; abdominal sternite III anteriomedially with a conspicuous depression marked by a thin rim. Constriction between abdominal segments III and IV deep. Abdominal segment IV very large, strongly recurved (
Whole body covered with dense mat of short, decumbent to suberect pubescent hairs; additionally, dorsal surfaces of body with abundant significantly longer suberect and erect hairs; such hairs also present on abdominal sterna III + IV, scapes (anterior faces of scapes with many hairs, posterior faces with fewer hairs) and legs (ventral faces of femora and tibiae with many hairs, dorsal faces with fewer hairs), the longest hairs on dorsal surface of body longer than the maximum dorsoventral diameter of metafemur. Mandibles striate; entire body densely punctate; on sides of pronotum punctures aligned in diffuse lines, appearing striate; punctures on antennae, legs, and abdominal segment IV finer than on rest of body; propodeal declivity shiny and at most superficially punctate; abdominal segments V–VII very superficially reticulate and shiny. Body color uniformly orange brown to reddish brown, vertex of head slightly darker, legs, antennal funiculus, and abdominal segments V–VII yellowish brown.
The species epithet is a patronym in honor of the German botanist Prof. Helge Bruelheide and his efforts in establishing and promoting the BEF-China project. All specimens of this species were collected on BEF-China field sites.
Most of the type series was collected during a leaf litter ant survey (
The variation in body size is within the normal limits of other
JAPAN, Fukuoka, Mt. Tachibana, 25-VII-1984, leg. S. Nomura (OKENT016137; OKENT016138; OKENT016139; OKENT016141; OKENT016142, all in
Volumetric raw data (in DICOM format), 3D rotation video (in .mp4 format, see Suppl. material
Still images from surface display volume renderings of 3D model of
This species is widely distributed, occurring from Japan (except Hokkaido) and South Korea to Vietnam. It has been recorded from Taiwan and the Chinese provinces Zhejiang and Hunan. Thus, we expect that it will be collected from the geographically intermediate provinces in the future. No direct biological observations from China are available, but the Japanese populations are comparatively well studied (
Still images from surface display volume renderings of 3D model of
In full-face view, head slightly longer than broad (
Mesosoma in profile slightly convex and slightly longer than maximum head length including mandibles (
Petiolar node in profile high, nodiform, with a straight and sloping anterior face, dorsum of node broadly rounded, posterior face half as long and steeper than anterior face; petiole in dorsal view longer than broad but apex of node clearly broader than long; ventral process moderately developed on anterior petiole, with a relatively indistinct rectangular projection.
In dorsal view abdominal segment III anteriorly much broader than petiole; its sides convex; abdominal sternite III anteriomedially with a conspicuous depression marked by a thin rim. Constriction between abdominal segments III and IV deep. Abdominal segment IV very large, recurved (
Whole body covered with dense mat of short, decumbent to suberect pubescent hairs; additionally, the dorsal surfaces of body interspersed with significantly longer suberect and erect hairs, such hairs also present on abdominal sterna III + IV, scapes (anterior faces of scapes with many hairs, posterior faces with single hairs), and legs (ventral faces of femora and tibiae with many hairs, dorsal faces with single hairs); the longest hairs on dorsal surface of body at most as long as the maximum dorsoventral diameter of metafemur. Mandibles striate; entire body including propodeal declivity densely punctate; on sides of pronotum punctures aligned in diffuse lines, appearing striate; punctures on antennae, legs, and abdominal segment IV finer than on rest of body, abdominal segments V–VII very superficially punctured and shiny. Body color uniformly orange brown to reddish brown, vertex of head slightly darker, frontal furrow conspicuously darker than surrounding cephalic dorsum, legs, antennal funiculus, and abdominal segments V–VII yellowish brown.
The species epithet is a patronym in honor of the Chinese botanist Prof. Keping Ma and his efforts in establishing the BEF-China project and promoting biodiversity research and nature conservation in China. All specimens of this species were collected in old-growth subtropical forest, an ecosystem Prof. Ma has investigated in detail.
Both specimens were collected in secondary mixed evergreen broadleaved forest of relatively advanced age, as indicated by the presence of large trees. The paratype was collected within the Gutianshan National Nature Reserve (
Apart from the small difference in body size (
This species is only known from the holotype that was collected in subtropical evergreen broadleaved forest at 2050 m asl. No direct observations of biology and natural history are available for
The unique hair patterns separate
Volumetric raw data (in DICOM format), 3D rotation videos (in .mp4 format, see Suppl. material
Still images from surface display volume renderings of 3D model of
Still images from surface display volume renderings of 3D model of
This species is only known from two locations at mid elevation in forests of southern and western Yunnan Province. The original description reported 45 workers in the type colony (
Even though at the beginning of this study we treated
This species was not mentioned in the revision of
Workers of this clade can be distinguished by a moderately squamiform petiolar node that narrows only little from base to apex (extremely squamiform in the Fiji archipelago,
The
JAPAN: Okinawa, Ishigaki Island, Mt. Omoto, 1-IV-1975, leg. M. Tanaka (
Volumetric raw data (in DICOM format), 3D rotation video (in .mp4 format, see Suppl. material
Still images from surface display volume renderings of 3D model of
This species is common from Japan (except Hokkaido) to Taiwan and usually collected in forests of relatively low elevation. It has also been reported from Yunnan Province in China. Thus, it is not unlikely that more records from the southern and eastern Chinese mainland will appear in the future if sampling effort is increased. No direct biological observations from China are available. In Japan, nests are typically found in deadwood in evergreen broadleaved forest (
According to
From
CHINA, Zhejiang Province, Gutianshan National Nature Reserve, ca. 30 km NW of Kaihua,
Volumetric raw data (in DICOM format), 3D rotation video (in .mp4 format, see Suppl. material
Still images from surface display volume renderings of 3D model of
The type locality is at ca. 1400 m asl in the Bà Nà hills close to Đà Nẵng city (referred to as Tourane in the original description), central Vietnam. The species is also known form Nangongshan Mountain, Mengla County, Yunnan Province (
This is a poorly known species. Since the single type specimen was not available for examination,
The only other
Worker of this clade can be separated from all other
This is an exclusively tropical clade with species occurring in Africa, Australia, Madagascar, the Mascarene Islands, Mesoamerica, and tropical southeast Asia. Eleven extant species are known, of which
Still images from surface display volume renderings of 3D model of
In full-face view, head slightly longer than broad (
Mesosoma in profile convex and longer than maximum head length including mandibles. Lower mesopleurae (katepisterna) with demarcated sutures, upper mesopleurae (anepisterna), and promesonotum with inconspicuous and very shallow sutures; lower mesopleurae inflated posteriorly; posterodorsal corners of propodeum with broad teeth that project over less than half of the propodeal lobes in profile, propodeal lobes strongly developed as broadly triangular teeth protruding dorsolaterally; propodeal declivity almost vertical, slightly inclined anteriorly; in posterodorsal view, sides of propodeum separated from declivity by lamellate margins; propodeal spiracle relatively small, located above mid height; in profile, opening ellipsoid and facing posteriorly. Legs comparatively long; all tibiae with a pectinate spur; calcar of strigil with a basal spine; pretarsal claws simple; arolia present.
Petiole in dorsal view longer than broad, sides consistently diverging posteriorly, anterior border with a thick margin that is distinctly angulate on each side; in profile, petiolar node relatively compressed dorsoventrally, its anterior face slightly sloping; dorsum of node relatively flat, weakly convex; ventral face inconspicuous with a thin lamella and no projection.
In dorsal view, abdominal segment III anteriorly much broader than petiole, its sides weakly convex; abdominal sternite III extended ventrally, its outline straight, anteriomedially with a conspicuous depression marked by a broad rim. Constriction between abdominal segments III and IV deep. Abdominal segment IV very large, very strongly recurved (abdominal sternum IV reduced and
Whole body covered with dense relatively short decumbent to erect hairs; additionally significantly longer suberect to erect hairs abundant on the whole body, including legs and scapes; such hairs also present on funicular joints, but shorter and relatively thicker; dense appressed to decumbent pubescence on the funiculus only; mandibles striate; head, mesosoma, petiole, and abdominal segment III foveolate with superimposed punctures and granules, the foveae relatively deep, large, and irregular; abdominal segment IV smooth and shiny, dorsally without sculpture, laterally superficially punctured; scapes and legs densely punctured. Body color uniformly dark ferruginous-brown, antennae, legs, and abdominal segments V–VII orange brown.
This species is named in honor of Dr. Shohei Suzuki (1979–2016), a Japanese marine biologist whose life was tragically lost in a diving accident while conducting coral reef research in Okinawa.
No direct observations of biology and natural history are available. The type specimen was collected from rain forest leaf litter. Like many other ant species occurring in the tropical rain forest of Xishuangbanna, the species probably also occurs in adjacent countries such as Laos or Thailand.
Maps of China (country is shown in dark grey with highlighted country and province borders) and South East Asia displaying known species distribution ranges (in green) of
Maps of China (country is shown in dark grey with highlighted country and province borders) and South East Asia displaying known species distribution ranges (in green) of
In
Since this species is known only from the holotype there is no available information about intraspecific variation.
As for most other regions in which
Recently,
With the exception of
Direct observations of ecology and natural history are very rare for Chinese
One problem encountered by
By applying
We thank the administration of the Gutianshan National Nature Reserve for granting research permissions for the forests under their management. The land of the BEF-China Main Experiment is owned by the Xingangshan Forestry Company, which is gratefully acknowledged for allowing research on their property. Merle Noack, Andreas Schuldt, Benoit Guénard, Benjamin Blanchard, and Masashi Yoshimura collected or provided specimens. We thank Adam Khalife for his assistance with processing the videos and Kenneth Dudley for the generation of the maps used in this study. The work of Michael Staab in China was funded by the German Research Foundation (DFG FOR 891/3, KL 1849/6-2), with travel support from the Sino German Center for Research Promotion (GZ 1146). Cong Liu, Francisco Hita Garcia, and Evan Economo were supported by subsidy funding from the Okinawa Institute of Science and Technology Graduate University.
Table S1
Data type: species data
Explanation note: Overview of specimens of non-Chinese
Figure S1.
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