Research Article |
Corresponding author: Li Ma ( maliwasps@aliyun.com ) Academic editor: Michael Ohl
© 2024 Yan Fu, Nawaz Haider Bashir, Qiang Li, Li Ma.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Fu Y, Bashir NH, Li Q, Ma L (2024) A review of Trypoxylon Latreille, 1796 (Hymenoptera, Crabronidae) of Southwest China with descriptions of two new species. Journal of Hymenoptera Research 97: 307-347. https://doi.org/10.3897/jhr.97.121279
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Two new species of the genus Trypoxylon (Hymenoptera: Crabronidae: Crabroninae: Trypoxylini) from Yunnan Province, China: T. aphelothoracicus Fu & Li, sp. nov. and T. ferrugineipes Fu & Li, sp. nov. are described and illustrated. The female of T. infoveatum Li & Li, 2007 is described for the first time. In addition, ten species of Trypoxylon are newly recorded from China: T. buddha Cameron, 1889, T. flavipes Tsuneki, 1979, T. fulvocollare Cameron, 1904, T. gampahae Tsuneki, 1981, T. imayoshii Yasumatsu, 1938, T. kandyianum Tsuneki, 1979, T. khasiae Cameron, 1904, T. nasale Tsuneki, 1979, T. pahangense Tsuneki, 1979, and T. pendleburyi Tsuneki, 1979. An updated key to Trypoxylon of Southwest China is provided.
Crabronidae, Identification key, new records, taxonomy
Southwest China, belonging to the main bioregions of Southeast Asia (Indochina), is recognized as one of the world’s 36 biodiversity hotspots and one of the regions with the richest and most threatened fauna worldwide. It’s located at the intersection of the Oriental and Palearctic regions, spanning subtropics and tropics and including plateau climate, tropical rainforest climate, and subtropical monsoon climate (
Southwest China, with an area of 2.5 million square kilometers, includes Yunnan Province, Guizhou Province, Sichuan Province, Chongqing Municipality, and Tibet Autonomous Region and is divided into three terrain units (Qinghai-Tibet Plateau, Yunnan-Guizhou Plateau, and Sichuan Basin), with the Hengduan Mountains, Yunnan-Guizhou Plateau, and Wushan Mountains regarded as the ‘Sky Islands of China’ and is a refuge for a wide range of flora and fauna, with an obvious vertical distribution of species (
Trypoxylon Latreille, 1796, has the widest distribution and most species (633 species and 84 subspecies) among the seven genera of Trypoxylini (Hymenoptera: Crabronidae: Crabroninae) (
Many authors studied the taxonomy of Trypoxylon.
In China, 55 species and nine subspecies of this genus are currently known, with 37 species and one subspecies found in southwest China (
The specimens examined are deposited in the Insect Collection of Yunnan Agricultural University, Kunming, China (YNAU). The specimens were observed and illustrated using an Olympus stereomicroscope (SZ Series) with an ocular micrometer. The photographs were taken with the VHX-5000 digital microscopic system and edited with Adobe Photoshop® 8.0. The descriptive terminology of morphological structures follows
AW apical width of the first flagellomere;
BW basal width of the apical flagellomere in male;
CV1, CV2 abscissa I of cubital vein, abscissa II of cubital vein;
F I, F II, F III, etc. the first, second and third flagellomere, etc.;
GL/ W ratio of gastral petiole length to apical width (dorsal view);
HL head length (frontal view);
HW head width (frontal view);
IOD interocular distance;
IODc minimum IOD at base of clypeus (frontal view);
IODv minimum IOD at vertex (dorsal view);
IODs ratio of IODv to IODc;
OOD ocellocular distance;
Od posterior ocellus diameter;
PD puncture diameter;
PIS puncture interspace;
POD postocellar distance;
R1 apical part of forewing vein RI beyond the meeting point with Rs;
TCV transverse cubital vein.
The frontal shield in some species has lateral bifurcation directed towards the eye incision; the upper area of the frontal shield is the area from the top to the base of the lateral bifurcation, and the lower area is from the base of the lateral bifurcation to the junction of the lateral carina in the frontal end.
Females
1 | Frons with shield-shaped enclosure; fore-wings with CV2 and TCV usually forming acute angle; dorsal and posterior area of propodeum with several conspicuous, transverse carinae | 2 |
– | Frons without shield-shaped enclosure; fore-wings with CV2 and TCV usually forming right or obtuse angle; dorsal and posterior area of propodeum without transverse carinae | 4 |
2 | Frontal shield discontinued, upper lateral carina broadly interrupted but dorsal carina clearly defined | Trypoxylon interruptum Tsuneki, 1978 |
– | Frontal shield complete, upper lateral carina and dorsal carina continued | 3 |
3 | Frontal shield with upper area subequal in length to lower area, at most 1.5 × as long as lower area, lateral carina of upper area curved; lateral surface of propodeum coriaceous, conspicuously obliquely rugose | Trypoxylon schmiedeknechtii Kohl, 1906 |
– | Frontal shield with upper area more than 1.5 × as long as lower area, lateral carina of upper area almost parallel; lateral surface of propodeum smooth medially and posteriorly, with inconspicuous oblique rugae anteriorly | Trypoxylon thaianum Tsunek, 1961 |
4 | Gastral terga I–III with apical fovea; pronotal collar narrow, with median tubercle | 5 |
– | Gastral terga I–III without apical fovea; pronotal collar broad, without median tubercle | 8 |
5 | Frons and mesoscutum with large punctures, PIS ≤ 0.5 × PD, PIS shiny; lateral surface of propodeum dull, with conspicuous oblique rugae; free margin of clypeus markedly concave laterally, with short and wide protrusion medially | Trypoxylon buddha Cameron, 1889 |
– | Frons and mesoscutum with fine punctures, PIS ≈ PD, PIS microscopically coriaceous; lateral surface of propodeum smooth, without rugae; free margin of clypeus straight or slightly convex laterally | 6 |
6 | Gastral tergum I without apicomedian fovea; legs black, at most partly brown | Trypoxylon bifoveatum Tsuneki, 1979 |
– | Gastral tergum I with apicomedian fovea; legs broadly yellow, only partly brown or black | 7 |
7 | Supraantennal tubercle with transverse subquadrate edge anteriorly; R1 equal to TCV, not reaching wing apex; gaster wholly black | Trypoxylon maculipes Tsuneki, 1979 |
– | Supraantennal tubercle rounded, without anterior transverse edge; R1 longer than TCV, almost reaching wing apex; gastral terga II–IV, base of gastral sternum III, gastral sternum IV ferruginous | Trypoxylon flavipes Tsuneki, 1979 |
8 | Gastral petiole clavate, gradually widening apically, as long as, or shorter than following two segments combined | 9 |
– | Gastral petiole flask-shaped, apical swelling rather abrupt, with parallel-sided stalk, longer than following two segments combined | 22 |
9 | Mandible thick, bidentate on inner margin near apex; head in frontal view quadrate, in dorsal view thick; median and lower frons roundly swollen | Trypoxylon gampahae Tsuneki, 1981 |
– | Mandible slender, without denticle on inner margin; head wider than long; median and lower frons not roundly swollen | 10 |
10 | Frontal furrow deeply impressed; legs slender and long (hind tibia about 1.25 × as long as HW, midtarsomere I longer than half HW), hind coxa more than three × apical width; propodeal dorsum long, more than 3.5 × as long as scutellum | Trypoxylon ferrugineipes Fu & Li, sp. nov. |
– | Frontal furrow very fine, inconspicuously impressed; legs thick and short (hind tibia about 0.93 × as long as HW, midtarsomere I shorter than half HW), hind coxa as long as, or shorter than twice apical width; propodeal dorsum short, shorter than 3.5 × as long as scutellum | 11 |
11 | Supraantennal tubercle with deep, longitudinal groove | 12 |
– | Supraantennal tubercle without deep, longitudinal groove | 13 |
12 | Sides of supraantennal tubercle with few rugae; gaster wholly black; IODs = 2:1 | Trypoxylon koreanum Tsuneki, 1956 |
– | Sides of supraantennal tubercle without rugae; gastral sterna II–III ferruginous brown, and apex of gastral petiole to sternum V yellow; IODs = 5:2 | Trypoxylon okinawanum Tsuneki, 1966 |
13 | Propodeal enclosure not delimited by boundary groove, densely covered with irregular, reticulate carinae; pronotal collar with black posterior band | 14 |
– | Propodeal enclosure delimited by more or less distinct U-shaped groove, surface smooth or covered with several transverse carinae medially; pronotal collar with light brown, translucent posterior band | 15 |
14 | Supraantennal tubercle conspicuously nasiform, with thick, longitudinal carina; clypeus with dense tiny punctures, free margin with small, rectangular protrusion medially, protruding area shallowly incised mesally | Trypoxylon fronticorne obliquum Tsuneki, 1981 |
– | Supraantennal tubercle low, with thin, longitudinal carina; clypeus with sparse large punctures, free margin with large, rectangular protrusion medially | Trypoxylon figulus (Linnaeus, 1758) |
15 | Supraantennal tubercle low, medial longitudinal carina thin | 16 |
– | Supraantennal tubercle highly nasiform, medial longitudinal carina thick | 20 |
16 | Gaster black, most of gastral sterna brownish | 17 |
– | Gaster more or less ferruginous | 18 |
17 | Free margin of clypeus with nearly triangular protrusion, bidentate mesally; side of propodeum with distinct lateral carina | Trypoxylon shimoyamai Tsuneki, 1958 |
– | Free margin of clypeus with inverted trapezoid protrusion medially; side of propodeum without lateral carina | Trypoxylon aphelothoracicus Fu & Li, sp. nov. |
18 | Free margin of clypeus rounded, without protrusion medially; U-shaped boundary groove on propodeal enclosure almost invisible and medial furrow shallow, surface smooth and shiny, without punctures or rugae | Trypoxylon truncatum Tsuneki, 1979 |
– | Free margin of clypeus with distinct protrusion medially; U-shaped boundary groove on propodeal enclosure and medial furrow clear and distinct, surface with conspicuous and dense rugae | 19 |
19 | Free margin of clypeus with distinct obtuse protrusion; supraclypeal area slightly narrow and long; gaster wholly ferruginous | Trypoxylon pahangense Tsuneki, 1979 |
– | Free margin of clypeus medially with distinctly inverted trapezoidal protrusion; supraclypeal area broad and short; gaster ferruginous from apex of petiole to apical gastral segment | Trypoxylon ferrugiabdominale Li & Li, 2007 |
20 | Free margin of clypeus conspicuously produced, with large semi-elliptic protrusion medially, as long as Od; gastral petiole and segments IV–VI black; legs wholly black | Trypoxylon clypeisinuatum Li & Li, 2010 |
– | Free margin of clypeus slightly produced medially, protrusion small, shorter than Od; gastral terga IV–VI ferruginous or with black maculae; legs partly ferruginous | 21 |
21 | Free margin of clypeus ferruginous, with semicircular protrusion, produced area shallowly incised mesally; gastral petiole broad and short, GL/ W = 2.7–3.0; gaster wholly ferruginous; all trochanters amber yellow | Trypoxylon nasale Tsuneki, 1979 |
– | Free margin of clypeus black, with two barely separated and round teeth medially; gastral petiole much slender, GL/ W = 3.5–3.9; gaster ferruginous from apex of petiole to apical segment; all trochanters black | Trypoxylon pendleburyi Tsuneki, 1979 |
22 | Gaster wholly or from apex of petiole to apical segment ferruginous | 23 |
– | Gaster black or middle part (from apex of petiole to segment III or IV or base of segments II–III) ferruginous | 25 |
23 | Gaster wholly ferruginous, petiole with black macula; supraantennal furrow absent; supraantennal tubercle low, with anterior transverse carina connected to antennal socket rim | Trypoxylon kandyianum Tsuneki, 1979 |
– | Gaster from apex of petiole to apical segment ferruginous; supraantennal furrow well developed; supraantennal tubercle without anterior transverse carina | 24 |
24 | Supraantennal tubercle attenuate apically, apex of supraantennal tubercle obliquely inclined, forming smooth and shiny area with large median hollow; margin of clypeus sinuate; antenna and legs mostly ferruginous; body length 11.9–12.5 mm | Trypoxylon khasiae Cameron, 1904 |
– | Supraantennal tubercle broaden apically, without anterior oblique flattened area; free margin of clypeus rounded; antenna and legs mostly black; body length 22.0 mm | Trypoxylon szechuen Tsuneki, 1981 |
25 | Mesoscutum microscopically coriaceous, with fine, dense punctures, PIS ≤ PD | 26 |
– | Mesoscutum smooth and shiny, with fine, scattered punctures, PIS > PD | 28 |
26 | Supraantennal tubercle without median carina, instead impressed line separated apex of supraantennal tubercle; pronotal collar thick | Trypoxylon bilobatum Tsuneki, 1961 |
– | Supraantennal tubercle with median carina; pronotal collar thin | 27 |
27 | Supraantennal tubercle low tuberiform; propodeal enclosure with clear U-shaped boundary groove and medial furrow; base of gastral segments II–IV and legs mostly ferruginous | Trypoxylon imayoshii Yasumatsu, 1938 |
– | Supraantennal tubercle highly nasiform, with deep groove medially; propodeal enclosure with vague U-shaped boundary groove, without medial furrow; gaster wholly and legs black | Trypoxylon infoveatum Li & Li, 2007 |
28 | Supraantennal furrow shallow; antennal socket rim anteriorly expanded into two separate cylinders; gaster wholly black | Trypoxylon takasago Tsuneki, 1966 |
– | Supraantennal furrow deep; antennal socket rim not expanded; gaster ferruginous medially | 29 |
29 | Setae on head and thorax golden; pronotal collar posteriorly and base of gastral segments II–III ferruginous; body length 22.2 mm | Trypoxylon fulvocollare Cameron, 1904 |
– | Setae on head and thorax silvery; pronotal collar black posteriorly, gaster ferruginous from apex of petiole to segment III or IV; body length 18.0–20.0 mm | 30 |
30 | Free margin of clypeus transversely produced mesally, slightly incised; gaster ferruginous from apex of petiole to segment III, darkly marked dorsally and ventrally; legs black; IODs = 1:1 | Trypoxylon orientale Cameron, 1904 |
– | Free margin of clypeus rounded mesally, not produced and incised; gaster ferruginous from apex of petiole to base of segment IV; legs black, with ferruginous spots; IODs varied | 31 |
31 | Side of propodeum with distinct lateral carina; all trochanters amber yellow; IODs = 1.5:1 | Trypoxylon errans Saussure, 1867 |
– | Side of propodeum without lateral carina; all trochanters amber black | 32 |
32 | Lateral tubercles of pronotum toothed; vertex conspicuously depressed; F I = 2.8–3.3 × AW; antenna mostly ferruginous beneath; body length 14.0–19.0 mm | Trypoxylon bicolor Smith, 1856 |
– | Lateral tubercles of pronotum triangular; vertex undepressed; F I = 2.0–2.5 × AW; antenna brown beneath; body length 10.0–19.0 mm | Trypoxylon petiolatum Smith, 1858 |
Males
1 | Frons with shield-shaped enclosure; fore-wings with CV2 and TCV usually forming acute angle; dorsal and posterior area of propodeum with several conspicuous, transverse carinae | 2 |
– | Frons without shield-shaped enclosure; fore-wings with CV2 and TCV usually forming right or obtuse angle; dorsal and posterior area of propodeum without transverse carinae | 4 |
2 | Frontal shield discontinued, upper lateral carina broadly interrupted but dorsal carina clearly defined; flagellomere III beneath with linear tyloids, flagellomere IV excavate beneath at base (apical flagellomere longer than two but shorter than three preceding articles combined) | Trypoxylon interruptum Tsuneki, 1978 |
– | Frontal shield complete, upper lateral carina and dorsal carina continued; flagellomeres not modified | 3 |
3 | Frontal shield with upper area as long as lower area, upper lateral carina curved; apical flagellomere longer than three but shorter than four preceding articles combined | Trypoxylon schmiedeknechtii Kohl, 1906 |
– | Frontal shield with upper area longer than lower area; apical flagellomere as long as three preceding articles combined | Trypoxylon thaianum Tsunek, 1961 |
4 | Gastral terga I–III with apical fovea; pronotal collar narrow, with median tubercle | 5 |
– | Gastral terga I–III without apical fovea; pronotal collar broad, without median tubercle | 8 |
5 | Frons and mesoscutum with large punctures, PIS ≤ 0.5 × PD, PIS shiny; lateral surface of propodeum dull, with conspicuous oblique rugae; lateral margin of clypeus slightly concave, with short, wide protrusion medially (penis valve subapically with narrow, curved hook on each side) | Trypoxylon buddha Cameron, 1889 |
– | Frons and mesoscutum with fine punctures, PIS ≈ PD, PIS microscopically coriaceous; lateral surface of propodeum smooth, without rugae; lateral margin of clypeus straight or slightly convex | 6 |
6 | Gastral tergum I without fovea; flagellomeres II–VI beneath with tyloids, flagellomeres VII–VIII excavate at base | Trypoxylon bifoveatum Tsuneki, 1979 |
– | Gastral tergum I with fovea; flagellomeres II–VI beneath without tyloids, flagellomeres V–VI excavate at base | 7 |
7 | Supraantennal tubercle subquadrate, surface nearly flat, including supraantennal furrow; R1 equal to TCV, not reaching wing apex; penis valve simple at apex | Trypoxylon maculipes Tsuneki, 1979 |
– | Supraantennal tubercle low, broad, roundly tuberiform, apical edge curved, not including supraantennal furrow; R1 longer than TCV, almost reaching apex of wing; penis valve subapically with narrow, curved hook on each side | Trypoxylon flavipes Tsuneki, 1979 |
8 | Gastral petiole clavate, as long as or shorter than segments II–III combined | 9 |
– | Gastral petiole flask-shaped, longer than segments II–III combined | 20 |
9 | Supraantennal tubercle with deep longitudinal groove | Trypoxylon koreanum Tsuneki, 1956 |
– | Supraantennal tubercle without deep longitudinal groove | 10 |
10 | Propodeal enclosure not delimited by boundary groove, densely covered with irregular, reticulate carinae; pronotal collar with black posterior band; penis valve with more or less pronounced preapical enlargement | 11 |
– | Propodeal enclosure delimited by more or less distinct U-shaped groove, surface smooth or covered with several transverse carinae medially; pronotal collar with light brown, translucent posterior band; penis valve without preapical enlargement | 12 |
11 | Supraantennal tubercle highly nasiform, with thick mid-longitudinal carina; flagellomeres without tyloids | Trypoxylon fronticorne obliquum Tsuneki, 1981 |
– | Supraantennal tubercle low, longitudinal carina thin; flagellomeres III–VIII beneath with tyloids | Trypoxylon figulus (Linnaeus, 1758) |
12 | Median and lower frons flat, without apical transverse carina and medial carina (flagellomeres I–XI beneath with tyloids, flagellomere IV excavate at base beneath) | Trypoxylon planifrons Tsuneki, 1977 |
– | Median and lower frons raised, with apical transverse carina or medial carina | 13 |
13 | Supraantennal tubercle highly nasiform, medial longitudinal carina thick | 14 |
– | Supraantennal tubercle low, medial longitudinal carina narrow | 17 |
14 | Antenna without tyloids | 15 |
– | Antenna with tyloids | 16 |
15 | Apical flagellomere as long as three preceding articles combined; gaster ferruginous, with vaguely outlined, black, regular-shaped band on gastral terga I–IV | Trypoxylon fenchihuense Tsuneki, 1967 |
– | Apical flagellomere as long as four preceding articles combined; gaster brown to black | Trypoxylon similichingi Li & Li, 2010 |
16 | Flagellomeres II–IV beneath with linear tyloids, apical flagellomere as long as, or shorter than five preceding articles combined | Trypoxylon clypeisinuatum Li & Li, 2010 |
– | Flagellomeres III–IV beneath with linear tyloids, apical flagellomere as long as four preceding articles combined | Trypoxylon pacificum Gussakovskij, 1932 |
17 | Apical flagellomere curved, conspicuously hollowed beneath (as long as three preceding articles combined); U-shaped boundary groove on propodeal enclosure almost invisible and medial furrow shallow, surface smooth and shiny, without punctures or rugae | Trypoxylon truncatum Tsuneki, 1979 |
– | Apical flagellomere not curved; U-shaped boundary groove on propodeal enclosure and medial furrow clear, distinct, surface with conspicuous dense rugae | 18 |
18 | Flagellomeres without tyloids and not excavate beneath (apical flagellomere as long as four preceding articles combined) | Trypoxylon ferrugiabdominale Li & Li, 2007 |
– | Flagellomeres with tyloids | 19 |
19 | Flagellomeres V–VI stoutly dentate beneath, apical flagellomere as long as two preceding articles combined | Trypoxylon shimoyamai Tsuneki, 1958 |
– | Flagellomeres not dentate beneath, flagellomeres III–VIII beneath with linear tyloids, apical flagellomere as long as three preceding articles combined | Trypoxylon kansitakum Tsuneki, 1971 |
20 | Apical flagellomere as long as or longer than four preceding articles combined | 21 |
– | Apical flagellomere shorter than four preceding articles combined | 23 |
21 | Flagellomere VIII excavate beneath at base, distinctly incrassate toward apex; supraantennal tubercle highly nasiform, with deep longitudinal groove at base | Trypoxylon infoveatum Li & Li, 2007 |
– | Flagellomeres unmodified; supraantennal tubercle tuberiform, without groove | 22 |
22 | Apical flagellomere longer than five preceding articles combined; setae on head and thorax silvery; gastral segments II–III ferruginous | Trypoxylon errans Saussure, 1867 |
– | Apical flagellomere as long as four preceding articles combined; setae on head and thorax golden; apex of gastral segments II–III ferruginous | Trypoxylon fulvocollare Cameron, 1904 |
23 | Mesoscutum distinctly microscopically coriaceous, superimposed with punctures, PIS ≈ PD, PIS coarse (apical flagellomere longer than two preceding articles combined; base of gastral segments II–IV ferruginous) | Trypoxylon imayoshii Yasumatsu, 1938 |
– | Mesoscutum without microsculpture, simply punctated, PIS > PD, PIS smooth and shiny | 24 |
24 | Supraantennal furrow shallow, antennal socket rim anteriorly expanded (supraantennal tubercle round, without transverse carina or band-like expansion at anterior margin; apical flagellomere longer than three preceding articles combined) | Trypoxylon takasago Tsuneki, 1966 |
– | Supraantennal furrow deep, antennal socket rim not expanded | 25 |
25 | Supraantennal tubercle attenuate apically, apex of supraantennal tubercle obliquely inclined, forming smooth and shiny area with large hollow mesally (apical flagellomere longer than two preceding articles combined and shorter than three preceding articles combined) | Trypoxylon khasiae Cameron, 1904 |
– | Supraantennal tubercle broaden apically, without anterior oblique flattened area | 26 |
26 | Side of propodeum with distinct lateral carina; clypeus conspicuously protruded medioapically; gaster from apex of petiole to segment III ferruginous laterodorsally, dark dorsally and ventrally | Trypoxylon orientale Cameron, 1904 |
– | Side of propodeum without lateral carina; clypeus round medioapically; gaster ferruginous from apex of petiole to base of gastral segment IV | 27 |
27 | Lateral tubercles of pronotum dentate; vertex conspicuously depressed; apical flagellomere in lateral view distinctly tapering | Trypoxylon bicolor Smith, 1856 |
– | Lateral tubercles of pronotum triangular; vertex undepressed; apical flagellomere in lateral view not tapering, slightly curved medially | Trypoxylon petiolatum Smith, 1858 |
Holotype : ♀: China, Yunnan Province, Jinghong City, Menghai County, Bulang Mountain, 21°37'35"N, 100°24'23"E, 1438 m., 20.VI–20.VII.2018, Li Ma project team (YNAU). Paratypes: 29♀♀: same locality as for holotype except: 20.VI–20.VII.2018 (10♀♀), 20.VII–15.VIII.2018 (3♀♀), 17.V–21.VI.2018 (2♀♀), 25.IV–17.V.2018 (3♀♀), 28.V–28.VI.2019 (10♀♀), 15.IV–27.V.2021 (1♀); 1♀, China, Yunnan, Jinghong City, Menghai County, Guanggang Village, Ancient tea forest, 21°49'15"N, 100°29'44"E, 1526 m, 20.VIII–16.IX.2018, coll. Li Ma project team (YNAU).
The species resembles T. minutum Tsuneki, 1979 and T. undatum Tsuneki, 1979 in lacking the lateral carina on the propodeum. It differs from both by the supraantennal tubercle with small U-shaped carina, transverse carina on both sides of apex, and with short, longitudinal carina mesally (in T. minutum the supraantennal tubercle is triangular, without transverse carina anteriorly and without middle carina; in T. undatum the supraantennal tubercle is low, with conspicuous, transverse carina anteriorly and thick, longitudinal carina mesally), free margin of clypeus with an inverted trapezoid projection (in T. minutum the free margin of clypeus is triangularly produced; in T. undatum the clypeal margin is wavy, without projection), gastral sterna II–IV black, gastral terga II–IV brown to black (in T. minutum gastral sterna II–IV are dark red, gastral terga II–IV are ferruginous; in T. undatum gastral sterna and terga II–IV are ferruginous, gastral terga II–III each with broad brown mark).
Female: Body length, 6.9–7.2 mm (Fig.
Head
: Head quadrate in frontal view (Fig.
Thorax
: Pronotum with deep, transverse furrow anteriorly, convex laterally, flattened anteriorly, pronotal collar narrow medially, enlarged towards side, without median tubercle, with distinct, translucent, posterior border; pronotal lobes rounded. Mesoscutum (Fig.
Gaster
: Gastral petiole (Fig.
Male. Unknown.
China (Yunnan).
The specific name is derived from two Greek words: apheles - (=smooth) and - thoracicus (= Latinized form of thorax), referring to the mesopleuron, metapleuron, and propodeal lateral surface smooth, and the side of propodeum without lateral carina in the female.
Holotype : ♀: China, Yunnan Province, Jinghong City, Menghai County, Bulang Mountain, 21°37'35"N, 100°24'23"E, ca 1438 m, 21.VI–20.VII.2018, Li Ma project team (YNAU). Paratype: 3♀♀: same date as holotype except: 28.V–28.VI.2019 (2♀♀), 13.VIII–15.IX.2020 (1♀).
The species resembles T. longipes Tsuneki, 1979 in having the legs markedly slender and elongate (hind tibia about 1.25 × as long as HW, midtarsomere I longer than half HW), free margin of clypeus wavy, supraantennal tubercle low and supraantennal furrow shallow. It differs by the IODs = 10:7 (in T. longipes the IODs = 10:4), gastral petiole slightly flask-shaped, GL/ W = 4.1 (in T. longipes gastral petiole distinctly flask-shaped, GL/ W = 5.6), gaster wholly ferruginous (in T. longipes gastral tergum V blackish). The species also resembles T. ambiguum Tsuneki, 1956 in the shape of the clypeal free margin and pronotal collar, but has a shallow supraantennal furrow (in T. ambiguum the supraantennal furrow is absent), IODs = 10:7 (in T. ambiguum IODs = 10:9), gastral petiole slightly clavate (in T. ambiguum gastral petiole flask-shaped), GL/ W = 4.1 (in T. ambiguum GL/ W = 5.0).
Female: Body length, 7.7–8.0 mm (Fig.
Head
: Head rounded in frontal view (Fig.
Thorax
: Pronotum with deep transverse furrow anteriorly, convex laterally, flattened anteriorly, pronotal collar narrow medially and enlarged towards side, with minute median tubercle, with distinct, translucent, posterior border; pronotal lobe rounded. Mesoscutum (Fig.
Gaster
: Gastral petiole (Fig.
Male. Unknown.
China (Yunnan).
The specific name is derived from two Latin words: ferrugineus - (= ferruginous) + - pes (= leg), referring to the legs partly ferrugineus in female.
Trypoxylon infoveatum Li & Li, 2007: 6.
6♀♀: China, Yunnan Province, Jinghong City, Menghai County, Bulang Mountain, 21°37'35"N, 100°24'23"E, ca 1438 m, 21.VI–20.VII.2018 (1♀), 25.IV–17.V.2018 (1♀), 19.IV–28.V.2019 (2♀♀), 28.V–28.VI.2019 (2♀♀), Li Ma project team (YNAU); 15♂♂: same data as for preceding: 25.IV–17.V.2018 (1♂), 17.V–21.VI.2018 (1♂), 21.VI–20.VII.2018 (1♂), 20.VII–15.VIII.2018 (1♂), 15.IX–16.X.2018 (2♂♂), 16.X–17.XI.2018 (2♂♂), 26.II–22.III.2019 (1♂), 22.III–19.IV.2019 (1♂), 28.VI–19.VII.20198 (3♂♂), 13.VIII–15.IX.2020 (1♂), 27.V–15.VI.2021 (1♂).
The species resembles T. koreanum Tsuneki, 1956 and T. koikense Tsuneki, 1956 in having the supraantennal tubercle highly nasiform, with deep groove medially. It differs from both by gastral petiole flask-shaped, longer than following two segments combined, GL/ W = 5.25 (in T. koreanum and T. koikense the gastral petiole is clavate, shorter than following two segments combined, GL/ W = 2.0–2.7), propodeal dorsum without mid furrow, obliquely carinae (in T. koreanum and T. koikense the propodeal dorsum with mid furrow, without oblique carinae), flagellomere VIII excavate beneath at base and distinctly incrassate toward apex in male (in T. koreanum flagellomeres unmodified in male; in T. koikense flagellomere VIII unmodified but flagellomere IV excavate beneath at base in male).
Female (first description of female): Body length, 7.5 mm (Fig.
Trypoxylon infoveatum Li & Li, 2007. ♀ (A, D, F, G, J, L); ♂ (B, C, E, H, I, K, M, N, O, P) A, B habitus (lateral view) C male antenna (lateral view) D, E head (frontal view) G, H thorax (dorsal view) F, I propodeum (dorsal view) J, K thorax (lateral view) L, M gastral segments I–III (dorsal view) N male sternum VIII O, P genitalia.
Head
: Head rounded in frontal view (Fig.
Thorax
: Pronotum with deep transverse furrow anteriorly, convex laterally, flattened anteriorly, pronotal collar narrow medially, enlarged towards side, with minute median tubercle, with distinct black posterior border; pronotal lobe rounded. Mesoscutum microscopically coriaceous (Fig.
Gaster
: Gastral petiole (Fig.
Male: Sculpture, setae, and body coloration (Fig.
China (Yunnan).
Trypoxylon buddha
Cameron, 1889: 118, 119;
Trypoxylon monstruosum
Tsuneki, 1974: 633, synonymized with Trypoxylon buddha by
Trypoxylon buddhae tarawakanum
Tsuneki, 1976: 92, synonymized with Trypoxylon buddha by
1♀: China, Yunnan Province, Jinghong City, Mengla County, Longmen Village, 21°16'46"N, 101°32'19"E, ca 923 m, 10.IV.2010, Rui Zhang (YNAU).
T. buddha resembles T. brevipenne de Saussure, 1867 in having large punctures on the frons and mesoscutum, metapleural keel conspicuously curved and hind coxae with tubercle in female. It differs by the apex of gastral terga I–III each with apicomedian fovea (in T. brevipenne the gastral terga I–III without fovea), free margin of clypeus with short, wide protrusion medially (in T. brevipenne free margin of clypeus with semicircular protrusion medially). The species also resembles T. maculipes Tsuneki, 1979 in sharing the apex of gastral terga I–III each with apicomedian fovea and in body colour, but the punctures on frons and mesoscutum are large (in T. maculipes punctures on the frons and mesoscutum are small), supraantennal tubercle with middle carina and anterior transverse carina (in T. maculipes supraantennal tubercle without middle carina and anterior transverse carina inconspicuous), free margin of clypeus with short, wide protrusion medially (in T. maculipes free margin of clypeus with bidentate protrusion medially).
Female (first record from China): Body length 9.5 mm (Fig.
China (Yunnan); India; Philippines; Sri Lanka.
Trypoxylon flavipes Tsuneki, 1979a: 3, 24, 1979b: 3, 8, 1980a: 4, 17, 1981a: 4, 13, 1981b: 100, 103, 1981d: 18, 1981f: 43.
2♀♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Rainforest, 21°91'37"N, 101°27'07"E, ca 606 m, 24.IV–31.V.2019, Yongsheng Pu (YNAU).
T. flavipes resembles T. buddha Cameron, 1889 and T. maculipes Tsuneki, 1979 in having the apex of gastral terga I–III each with apicomedian fovea and head sub-quadrate in frontal view. It differs from both by the anterior edge of supraantennal tubercle rounded (in T. buddha supraantennal tubercle with conspicuous transverse carina anteriorly; in T. maculipes the anterior edge of supraantennal tubercle transverse), the gaster and legs more or less ferruginous (in T. buddha and T. maculipes the gaster and legs wholly black), punctures on the frons and mesoscutum are small (in T. buddha punctures on the frons and mesoscutum are large), free margin of clypeus with bidentate protrusion medially (in T. buddha free margin of clypeus with short, wide protrusion medially), the penis valve subapically with narrow, curved hook on each side (in T. maculipes the penis valve is simple at apex).
Female (first record from China): Body length 7.7–7.8 mm (Fig.
Australia; Borneo; China (Yunnan); India; Laos; Moluccas; New Guinea; Pacific Islands; Philippines; Sri Lanka; Sulawesi.
Trypoxylon fulvocollare
Cameron, 1904: 217;
1♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Rainforest, 21°91'37"N, 101°27'07"E, ca 606 m, 19.VI–13.VII.2021, Yongsheng Pu (YNAU).
T. fulvocollare resembles T. taiwanum Tsuneki, 1967 and T. atricorne Tsuneki, 1979 in having the supraantennal tubercle low, with thin mid-longitudinal carina, without anterior carina, the antennal socket rim tricarinate, the shape of pronotal collar and punctures on the frons and mesoscutum fine and sparse. It differs from both by the body covered with golden setae (in T. taiwanum and T. atricorne the setae are silvery), the flagellomeres I–II beneath and pronotal collar posteriorly yellow (in T. taiwanum the pronotal collar posteriorly black to light brown; in T. atricorne the flagellomeres I–II black and pronotal collar posteriorly are black to light brown), the base of gastral segments II–III are ferruginous (in T. taiwanum and T. atricorne the gaster is ferruginous from apex of petiole to segment III or IV), OOD: POD = 2: 3 (in T. taiwanum OOD: POD = 1: 3 and in T. atricorne OOD: POD = 1: 2), free margin of clypeus rounded (in T. taiwanum free margin of clypeus is conspicuously rounded; in T. atricorne free margin of clypeus is rounded and shallowly incised mesally).
Female (first record from China): Body length 21.2 mm (Fig.
Borneo; China (Yunnan); Indonesia; Java; Lesser Sunda Islands; Malaysia; Moluccas; Philippines; Sulawesi; Sumatra.
Trypoxylon gampahae Tsuneki, 1981d: 5, 19.
1♀: China, Yunnan Province, Jinghong City, Menghai County, Bulang Mountain, 21°37'35"N, 100°24'23"E, ca 1438 m, 27.V–15.VI.2021, Yongsheng Pu (YNAU).
T. gampahae resembles T. mandibulatum Richards, 1933 and T. pygmaeum Cameron, 1900 in having the mandible bidentate on inner margin near apex, median and lower frons roundly swollen and head sub-quadrate in frontal view. It differs from both by punctures on the frons and mesoscutum are fine and sparse (in T. mandibulatum punctures on the frons and mesoscutum somewhat are large and conspicuous), the frons in lateral view is highly raised and inclined to antennal socket rim anteriorly (in T. pygmaeum the frons in lateral view is inconspicuously raised and almost flat anteriorly), the free margin of clypeus is conspicuously produced and with bidentate protrusion medially (in T. mandibulatum the clypeal free margin is inconspicuously produced and with truncate protrusion medially; in T. pygmaeum the free margin of clypeus is inconspicuously produced and slightly wavied).
Female (first record from China): Body length 7.5 mm (Fig.
China (Yunnan); Sri Lanka.
Trypoxylon imayoshii
Yasumatsu, 1938: 451, 453;
1♀1♂: China, Guizhou Province, Zunyi City, Leigong Mountain, 27°52'06"N, 106°56'53"E, 1184 m, 19.VII.2019 (1♀), 15.VII.2019 (1♂), Yan Zhang (YNAU). 2♀♀3♂♂: China, Guizhou Province, Zunyi City, Dabanshui Forest Park, 27°42'18"N, 106°51'15"E, ca 1001 m, 10. VII. 2011, Feng Dongdong (YNAU).
T. imayoshii resembles T. ambiguum Tsuneki, 1956 in having a similar shape of the supraantennal tubercle, the pronotal collar and gastral petiole, punctures on the frons and mesoscutum are fine and sparse. It differs by the free margin of clypeus is conspicuously produced and with slightly bidentate protrusion medially (in T. ambiguum the margin of clypeus is inconspicuously produced and slightly waved), the frontal furrow is conspicuously impressed, the surface gently inclined towards middle (in T. ambiguum the frontal furrow is inconspicuous, only conspicuous before anterior ocellus), the flagellomere XI longer than two preceding articles combined in male (in T. ambiguum the flagellomere XI shorter than two preceding articles), the apex of sternum VIII rounded in male (in T. ambiguum the apex of sternum VIII incised mesally in male). The species also resembles T. infoveatum Li & Li, 2007 in having the shape of pronotal collar and gastral petiole, punctures on the frons and mesoscutum are fine and sparse, but the supraantennal tubercle is low (in T. infoveatum the supraantennal tubercle is highly nasiform), the gaster is more or less ferruginous (in T. infoveatum the gaster is wholly black), the flagellomeres are unmodified in male (in T. infoveatum the flagellomere VIII excavate beneath at base and markedly incrassate toward apex).
Female (first record from China): Body length 8.0–9.7 mm (Fig.
Trypoxylon imayoshii Yasumatsu, 1938. ♀ (A, D, F, G, J, L); ♂ (B, C, E, H, I, K, M, N, O, P) A, B habitus (lateral view) C male antenna (lateral view) D, E head (frontal view) G, H thorax (dorsal view) F, I propodeum (dorsal view) J, K thorax (lateral view) L, M gastral segments I–III (dorsal view) N male sternum VIII O, P genitalia.
Male. Body length 7.0–9.6 mm (Fig.
China (Fujian, Guangdong, Guangxi, Shandong, Yunnan, Zhejiang); Japan; Korea; Russia.
Trypoxylon kandyianum Tsuneki, 1979b: 4, 17, 1981d: 19.
1♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Rainforest, 21°91'37"N, 101°27'07"E, ca 606 m, 24.IV–31.V.2019, Yongsheng Pu (YNAU); 1♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Sandalwood and pomelo mixed forest, 27°90'04"N, 106°27'21"E, ca 508 m, 25.IX–25.X.2019, Yongsheng Pu (YNAU).
The species resembles T. formosicola Strand, 1922 in having the supraantennal tubercle is low, but differs by rounded free margin of clypeus (in T. formosicola free margin of clypeus have short, wide protrusion), the scape and pedicel beneath and gaster are wholly ferruginous (in T. formosicola the scape and pedicel beneath and gaster are wholly black). The species also resembles T. gracilescens F. Smith, 1860 in having the free margin of clypeus rounded. It differs by the supraantennal furrow is absent (in T. gracilescens the supraantennal furrow is deep), the side of the propodeum have conspicuous lateral carina (in T. gracilescens the lateral carina is inconspicuous, almost lacking), the supraantennal tubercle is low, with anterior transverse carina connected to antennal socket rim (in T. gracilescens the medio-apical area of supraantennal tubercle is obliquely flattened into smooth, shiny and round area, not connected to the antennal socket rim).
Female (first record from China): Body length 11.9–12.5 mm (Fig.
China (Yunnan); Sri Lanka.
Trypoxylon khasiae
Cameron, 1904d: 218;
3♀♀: China, Yunnan Province, Honghe Prefecture, Hekou County, Nanxi Town, 22°37'32"N, 103°56'53"E, ca 121 m, 6.VIII.2016 (2♀♀), 28.X.2016 (1♀), Hesheng Wang (YNAU); 3♀♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Rubber forest, 21°92'26"N, 101°26'50"E, ca 543 m, 20.VIII–18.IX.2018, Li Ma project team (YNAU); 1♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Rainforest, 21°91'37"N, 101°27'07"E, ca 606 m, 20.VIII–18.IX.2018, Li Ma project team (YNAU); 1♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Sandalwood and pomelo mixed forest, 27°90'04"N, 106°27'21"E, ca 508 m, 26.VIII–26.IX.2019, Li Ma project team (YNAU).
T. khasiae resembles T. varipilosum Cameron, 1901 and T. hyperorientale Strand, 1922 in having the medio-apical area of supraantennal tubercle obliquely flattened into smooth, shiny and round area, the shape of pronotal collar, and a smooth mesoscutum. It differs from both by the body with silvery setae (in T. varipilosum the setae brassy), oblique area of supraantennal tubercle carrying fovea on it (in T. varipilosum oblique area is flat), IODs = 10:5.6 (in T. varipilosum IODs = 10:9.0), the free margin of clypeus is rounded out, with two notches medially (in T. hyperorientale the margin of clypeus is rounded, slightly incised mesally), flagellomeres beneath and gaster from apex of petiole to apical gastral segment are ferruginous (in T. hyperorientale the flagellomeres and gaster are wholly black).
Female (first record from China): Body length 11.9–12.5 mm (Fig.
China (Yunnan); India; Indonesia; Laos; Thailand.
Trypoxylon nasutum Tsuneki, 1979a: 5, 37.
Trypoxylon nasale Tsuneki, 1980a: 2. Substitute name for Trypoxylon nasutum Tsuneki, 1979.
Trypoxylon minahime Tsuneki, 1992: 54. Unnecessary substitute name for Trypoxylon nasutum Tsuneki, 1979.
3♀♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Rainforest, 21°91'37"N, 101°27'07"E, ca 606 m, 24.IV–31.V.2019, Yongsheng Pu (YNAU); 3♀♀: China, Yunnan Province, Jinghong City, Mengla County, Xishuangbanna Tropical Botanical Garden, Rubber forest, 21°92'26"N, 101°26'50"E, ca 543 m, 15.V–18.VI.2018, Lin Zhao (YNAU).
T. nasale resembles T. sauteri Tsuneki, 1981 and T. clypeisinuatum T. Li and Q. Li, 2010 in having the supraantennal tubercle is high nasiform, with anterior transverse carina connected to antennal socket rim. It differs from both by IODs = 10:8 (in T. sauteri IODs = 10:3.4), the fore- and midlegs except base of coxa and gaster are ferruginous (in T. sauteri the gastral segment I, V and VI are black; in T. clypeisinuatum the legs are wholly black, the gastral segment I, V and VI are black), free margin of clypeus have semicircular protrusion, the protrusionis shallowly incised mesally (in T. sauteri the margin of clypeus is slightly semicircularly produced; in T. clypeisinuatum the margin of clypeus is conspicuously semicircularly produced).
Female (first record from China): Body length 8.7–9.2 mm (Fig.
China (Yunnan); Malaysia.
Trypoxylon pahangense Tsuneki, 1979a: 6, 51, 1981d: 26, 38.
1♀: China, Yunnan, Jinghong City, Menghai County, Guanggang Village, Ancient tea forest, 21°49'15"N, 100°29'44"E, ca 1526 m, 20.VIII–16.IX.2018, coll. Li Ma project team (YNAU);1♀: China, Yunnan Province, Jinghong City, Menghai County, Bulang Mountain, 21°37'35"N, 100°24'23"E, ca 1438 m, 17.V–20.VI.2018, Li Ma project team (YNAU); 13♀♀: same data as for preceding: 20.VI–20.VII.2018 (2♀♀), 20.VII–16.VVIII.2018 (3♀♀), 16.VIII–14.IX.2018 (3♀♀), 28.V–28.VI.2019 (1♀), 28.VI–19.VII.2019 (1♀), 19.VII–21.VIII.2019 (1♀), 21.VIII–20.IX.2019 (1♀), 10.VII–13.VIII.2020 (1♀).
T. pahangense resembles T. truncatum Tsuneki, 1979 and T. brunneimaculatum T. Li and Q. Li, 2007 in having the supraantennal tubercle is low, with anterior transverse carina connected to antennal socket rim. It differs from both by the free margin of clypeus have conspicuously obtuse-shaped protrusion (in T. truncatum the margin of clypeus is rounded, without projection; in T. brunneimaculatum the clypeal free margin is slightly semicircularly produced), the gaster is wholly ferruginous (in T. truncatum the gastral petiole is black; in T. brunneimaculatum the gastral segment I, V and VI are black).
Female (first record from China): Body length 7.7–10.1 mm (Fig.
China (Yunnan); Malaysia.
Trypoxylon pendleburyi Tsuneki, 1979a: 5, 36.
1♀, China, Yunnan, Jinghong City, Menghai County, Guanggang Village, farmland, 21°49'50"N, 100°28'20"E, ca 1229 m, 16.IX–19.X.2018, coll. Li Ma project team (YNAU); 9♀♀: same data as for preceding: 28.V–28.VI.2019 (1♀), 20.VII–23.VIII.2019 (2♀♀), 23.X–24.XI.2019 (2♀♀), 13.I–15.II.2021 (4♀♀).
T. pendleburyi resembles T. nasale Tsuneki, 1979 and T. clypeisinuatum T. Li and Q. Li, 2010 in having the supraantennal tubercle is highly nasiform, with anterior transverse carina connected to antennal socket rim. It differs from both by free margin of clypeus have two barely separated and round teeth medially (in T. nasale the margin of clypeus have semicircular protrusion, the protrusion is shallowly incised mesally; in T. clypeisinuatum the margin of clypeus is conspicuously produced and with semicircular protrusion), the all trochanter are black, the gaster from apex of petiole to end is ferruginous (in T. nasale the all trochanter and gaster are wholly ferruginous; in T. clypeisinuatum the gastral segment I, V and VI are black), the gastral petiole is much slender (in T. nasale the gastral petiole is broad and short).
Female (first record from China): Body length 7.5–9.2 mm (Fig.
Australia; Borneo; China (Yunnan); India; Laos; Moluccas; New Guinea; Pacific Islands; Sri Lanka; Sulawesi.
Species from southwest China are found in both the Oriental and Palearctic regions, highlighting the richness and uniqueness of the region’s biodiversity and reflecting the complexity and diversity of the region’s natural environment. This study lays the foundation for further research on the relationship between climate change, environmental heterogeneity, and the diversity of sphecid wasps in southwest China.
Southeast Asia has the greatest diversity of the genus Trypoxylon, with Indonesia and Malaysia harboring the highest species diversity (
The endemic and newly recorded species of China are mainly distributed in southern China, which is part of the Indochina bioregions. The discovery of new species and the revision of existing ones in this region will provide new insights into the biodiversity and biogeographic distribution of Indochina. Additionally, it will provide a basis for further research on the origin, diffusion paths, and historical evolution processes of biological species across various bioregions.
Prospect. On the basis of morphological classification studies and the analysis of insect gene sequences by molecular biology methods, insect species can be identified and classified more accurately, thereby solving the problem of some species being very similar in morphology and difficult to distinguish (
We would like to thank the editor and two reviewers for suggestions that helped improve this article. We express our heartfelt thanks to Wojciech J. Pulawski (California Academy of Sciences, California) for providing us with many valuable references. We are also grateful to Dr. Jing-xian Liu (South China Agricultural University, Guangzhou, China), Yong-sheng Pu, Yu Tao, and Ling Zhao (Yunnan Agriculture University, Kunming, China) for helping us collect specimens. This work was supported by the National Natural Science Foundation of China under Grant number 32270485, the Agricultural Basic Research joint project of Yunnan Province under Grant number 202101BD070001-004 and the Yunnan Provincial Department of Science and Technology “Yunnan Talent Program” plan (202303AM140032).