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The Diversity of Hornets in the Genus Vespa (Hymenoptera: Vespidae;

Vespinae), Their Importance and Interceptions in the United States

Article in Insect Systematics and Diversity · May 2020

DOI: 10.1093/isd/ixaa006

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 Insect Systematics and Diversity, (2020) 4(3): 2; 1–27
doi: 10.1093/isd/ixaa006
Research
Taxonomy

The Diversity of Hornets in the Genus Vespa

(Hymenoptera: Vespidae; Vespinae), Their Importance
and Interceptions in the United States

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Allan H. Smith-Pardo,1,4 James M. Carpenter,2 and Lynn Kimsey3
1
USDA-APHIS-PPQ, Science and Technology (S&T), Sacramento, CA, 2Department of Invertebrate Zoology, American Museum
of Natural History, New York, NY, 3Bohart Museum of Entomology, University of California, Davis, Davis, CA, and 4Corresponding
author, e-mail: [email protected]

Subject Editor: Heather Hines

Received 20 December, 2019; Editorial decision 11 March, 2020

Abstract
Hornets in the genus Vespa (Vespidae, Vespinae) are social wasps. They are primarily predators of other in-
sects, and some species are known to attack and feed on honeybees (Apis mellifera L.), which makes them a
serious threat to apiculture. Hornet species identification can be sometimes difficult because of the amount of
intraspecific color and size variation. This has resulted in many species-level synonyms, scattered literature,
and taxonomic keys only useful for local populations. We present a key to the world species, information on
each species, as well as those intercepted at United States Ports of Entry during the last decade. Images of all
the species and some of the subspecies previously described are also included.

Resumen
Los avispones (Vespidae: Vespinae: Vespa) son avispas sociales, depredadoras de otros insectos y algunas
de las especies muestran cierta preferencia por abejas, incluyendo las abejas melíferas (Apis mellifera L.)
convirtiéndose en una amenaza para la apicultura. La identificación de las especies puede ser difícil dado
que muchas especies muestran una gran variación inter-especifica que ha tenido como resultado multitud
de sinonimias en el género y además, porque la literatura en el género es dispersa y las claves taxonómicas
solo funcionan para poblaciones locales. En este trabajo, presentamos una clave para las especies del mundo,
información para cada especie así como de las especies interceptadas en puertos de entrada a los Estados
Unidos durante la última década. Se incluyen además imágenes para todas las especies así como para algunas
de las subespecies previamente descritas.

Key words: invasive species, quarantine significance, pest, diversity, social wasp

Hornets in the genus Vespa are large, predatory, eusocial wasps na- et al. (2014). These subspecies were all synonymized by Carpenter
tive to Europe and Asia. They prey on a wide diversity of insects, and Kojima (1997).
but several species are predatory on honeybees. Vespa nests can be Beggs et al. (2011) assessed the distribution, abundance, im-
physically large, with over 1,000 workers, but usually with hun- pact, and management of invasive species of Vespidae worldwide.
dreds of workers (Archer 2008). Nests can be aerial, attached to In the case of the hornets, several Vespa species have been intro-
tree branches or in shrubs, in crevices, under eaves or underground duced outside their native ranges, including V. crabro Linnaeus,
depending on the species. Depending on the latitude, nests can be 1758 (Hymenoptera: Vespidae) from Europe into the United States
either annual, started by a new queen every spring, or perennial, (Bequaert 1932, Shaw and Weidhaas 1956); V. velutina Lepeletier,
where young queens take over from old ones (Matsuura and Yamane 1836 (Hymenoptera: Vespidae) from Asia into Europe (Monceau
1990). Colonies in warm tropical climates tend to be perennial. et al. 2014); and V. tropica Linnaeus, 1758 (Hymenoptera: Vespidae)
These wasps exhibit a range of color patterns that vary geo- from continental Asia into the island of Guam (Anonymous 2016).
graphically. Previous authors have named numerous subspecies There are also records of other species collected outside their ranges
based solely on color patterns, but Vespa color patterns tend to without becoming established (see Kimsey and Carpenter 2012 for
grade from one region to the next Nguyen et al. (2006), Perrard North America).

© The Author(s) 2020. Published by Oxford University Press on behalf of Entomological Society of America. 1
All rights reserved. For permissions, please e-mail: [email protected].
2 Insect Systematics and Diversity, 2020, Vol. 4, No. 3

Some of the introduced Vespa species have shown to be environ- Specimens used for the images were from the American Museum
mental stressors as predators of already declining populations of native of Natural History (AMNH), New York, NY (C. Lebeau) and from
insects, spiders, and honeybees (Shah and Shah 1991, Abrol 1994, Choi the Bohart Museum of Entomology (BME), University of California,
et al. 2011, among others). They can also be competitors for food and Davis (S. Heydon). Images were captured and edited using a Nikon
nesting sites of native wasps (Cini et al. 2018), vectors of parasites or SMZ18 System with a Nikon DS-fi2 camera, the stacking software,
diseases that can affect honey bees and native wasps (Choi et al. 2011), Helicon Focus 6.0, and Photoshop Elements 12.
and impact human safety because of their aggressiveness and fatal reac- Type repositories are given in parentheses at the end of species entries
tions to their venom (Nguyen et al. 2010, Kularatne et al. 2014). in synonym lists. These include BASEL = Naturhistorisches Museum,
The invasion of V. velutina into Europe typifies these prob- Basel, Switzerland; BERLIN = Museum für Naturkunde, Berlin,
lems. These hornets prey on the domestic honey bee [Apis mellifera Germany; BUDAPEST = Zoological Department, Hungarian Natural
(Linnaeus) (Hymenoptera: Apidae)], disrupt the ecological role of History Museum, Budapest, Hungary; CALCUTTA = Zoological

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honeybees as pollinators (Villemant et al. 2006a, b; Monceau et al. Survey of India, Calcutta; CAMBRIDGE = Museum of
2014), have potentially altered local biodiversity (Fedele et al. 2019), Comparative Zoology, Harvard University, Cambridge,
and are potentially deadly to people allergic to their stings. Massachusetts, USA; COPEHAGEN = Statens Naturhistoriske
In Europe, the native hornet, V. crabro, is protected in some re- Museum, Copenhagen, Denmark; FUKUOKA = Kitakyushu
gions, such as in Germany (Federal Species Protection Ordinance- Museum of Natural History and Human History, Fukuoka,
BArtSchV/Federal Nature Conservation Act-BNatSchG). The Japan; GENOA = Museo Civico di Storia Naturale di Genova,
introduced V. velutina may compete for food resources with Genoa, Italy; KAGOSHIMA = Reimeikan-Kagoshima History
V. crabro, which is a more generalist predator (Shaw and Weidhaas Museum, Kagoshima, Japan; KUNMING = Kunming Natural
1956, Baracchi et al. 2010). According to Monceau et al. (2014), History Museum of Zoology, Kunming, Yunnan, China;
some beekeepers have reported increased V. crabro predation on LEIDEN = Rijksmuseum van Natuurlijke Historie (now called
honeybees since the introduction of V. velutina in Europe. the Nederlands Centrum voor Biodiversiteit Naturalis), Leiden,
This species was introduced into New York State in the mid- Netherlands; LOGAN = The American Entomological Institute,
1800s (de Saussure 1898). Since then, it has since spread throughout Logan, Utah, USA; LONDON-LS = Linnaean Society, London,
eastern North America (Kimsey & Carpenter 2012). Vespa tropica England; LONDON-NHM = The Natural History Museum, London,
was first found on Guam in 2016 (Anonymous 2016). Since then, it England; MUNICH = Zoologische Staatssammlung München,
has become so widespread on the island it is no longer under quar- Munich, Germany; OXFORD = Hope Entomological Collections,
antine, even though problems have arisen because it is aggressive and Oxford University Museum, Oxford, England; PARIS = Museum
may nest close to human settlements. National d’Histoire Naturelle, Laboratoire d’Entomologie,
Studies by Blanchard et al. (2008) and Yañez et al. (2012) had Paris, France; SAPPORO = Entomological Institute, Hokkaido
shown the potential transmission of the Israeli Acute Paralysis Virus University, Sapporo, Japan; ST. PETERSBURG = Zoological
(IAPV), one of the causes of Colony Collapse Disorder in honeybees Museum of the Zoological Institute, Saint Petersburg, Russia;
(Cox-Foster et al. 2007) between honeybees and their non-Apis TAICHUNG = Taiwan Agricultural Research Institute, Taichung,
predators, such as hornets. The possibility of transmission from hor- Taiwan; TOKYO = National Museum of Nature and Science,
nets to honeybees or other native species has yet to be determined. Tokyo, Japan; TURIN = Museo Regionale di Scienze Naturali,
The introduction of alien hornet species into the United States is Turin, Italy; UPPSALA = Uppsala universitet Evolutionsmuseet,
a real risk and having a practical synthesis of the current knowledge Uppsala, Sweden; VENICE = Museo di Storia Naturale di Venezia
on the basic biology and identification of the world’s hornets would Venice, Italy; VIENNA = Naturhistorisches Museum, Vienna,
greatly increase our capacity to identify and enhance the detection of Austria; WASHINGTON = Smithsonian National Museum of
potentially invasive species of hornets or to monitor their dispersion Natural History, Washington, DC, United States.
if they are introduced.
In this study, we provide information on the species diversity of
hornets, taxonomic keys and visual aids for the identification of the Genus Vespa Linnaeus
different species. Vespa Linnaeus, 1758: 343. Type species: ‘Vespa crabro Fab’. (=
Vespa crabro Linnaeus, 1758); designated by Latreille, 1810: 438.
Macrovespa Dalla Torre, 1904: 64 (subgenus of Vespa Linnaeus).
Materials and Methods Type species Vespa crabro Linnaeus, 1758. Designated by
We performed an Ad hoc/ unlimited search for interceptions of spe- Bequaert, 1930: 64.
cies of Vespinae (Vespa and Vespula Thomson) at Ports of Entry Nyctovespa van der Vecht, 1959: 210 (subgenus of Vespa Linnaeus).
(PE) to the United States using the USDA-APHIS, Plant Protection Type species: Vespa binghami du Buysson, 1905, original
and Quarantine (PPQ) databases AQAS (Agricultural Quarantine designation.
Activity Systems) and ARM (Agricultural Risk Management System)
going back to 2010. After the search, the data were processed, fil- Diagnosis. The subfamily Vespinae can be distinguished from other
tered and the total of interceptions for hornets (genus Vespa) was Vespidae by the hind wings lacking an anal lobe (Fig. 1a); forewing
determined for the years 2010 to 2018. In addition, some informa- recurrent veins ending in the same submarginal cell (Fig. 1b), mar-
tion from interception records, such as origin and means of transpor- ginal cell narrowly pointed along costal vein (not extending away
tation, is given in the results and the discussion. from it) (Fig. 1b); pronotal lobe separated from tegula by a distance
Taxonomic keys provided below were constructed in part based equal or less than its length (Fig. 1c); tarsal claws simple, not bifid
on previous keys by Archer (1989, 2012). A revised, fully illustrated (Fig. 1d), without parategula (Fig. 1e), and metasoma sessile with
key was needed to clarify and simplify diagnostic characters used in first tergum truncate (Fig.1f).
previous keys. In addition to the keys, we provide, for the first time, Species of the genus Vespa can be distinguished from other ves-
high-resolution images of these features. pine genera by having both the pronotal and the pretegular carinae
Insect Systematics and Diversity, 2020, Vol. 4, No. 3 3

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Figs. 1–4. Vespa morphology. Fig. 1 (a) Hindwing. (b) Forewing apex. (c) Lateral view of pronotal lobe. (e) Hindtarsal claws. (f) Thoracic dorsum. (g) Lateral view
of metasomal segment 1. Fig. 2. (a) Lateral view of pronotal lobe. (b) Dorsal view of vertex, i=distance from hind ocellus to posterior head margin, ii=distance
between lateral ocellus and eye. (c) Lateral view of metasomal segment 1. (d) Costal margin of forewing apex. (e) Dorsal view of vertex. (c) Front view of clypeus
and mandibles. Fig. 3. (a, b) Lateral view of head. (c, d) Lateral view of metasoma. (3a, 3c) V. soror. Fig. 4. (a, b) Lateral view of metasomal tergum 2. (c, d) Lateral
view of pronotal carina. (e, f) Front view of clypeus. Figs. 1a-g, 2a, 2b, 3b, 3d, V. crabro. Fig. 2c, V. binghami. Figs. 2d, 2e, V. analis. Fig. 2f, V. bicolor. Figs. 4a, 4e,
V. fervida. Fig. 4b, V. multimaculata. Figs. 4c, 4f, V. luctuosa. Fig. 4d, V. affinis.

(Fig. 2a), head vertex with the distance from the posterior ocellus Keys to the Queens and Workers of the
to the posterior margin of vertex more than twice the distance be- Species of Vespa of the World
tween the posterior ocelli and the compound eye (Fig. 2b), the basal
1 Posterior ocelli about as close to compound eye as to each other
metasomal segment is anteriorly rounded (Fig. 1f), and the length
(Fig. 2c) ������������������������������������������������� binghami du Buysson
of the forewing prestigma is three times or more the length of the
- Posterior ocelli closer to each other than to compound eye
pterostigma (Fig. 1b).
(Fig. 2d)����������������������������������������������������������������������������������2
4 Insect Systematics and Diversity, 2020, Vol. 4, No. 3

2 Apical margin of clypeus with medial tooth between laterally usually reddish brown (Fig. 10d), or if metasomal tergum is
produced margins (Fig. 2e)������������������������������ analis Fabricius mostly black then the vertex is also black����� affinis (Linnaeus)
- Apical margin of clypeus without medial tooth between lat- 14 Clypeus strongly bulging medially, strongly convex in side view
erally produced margins (Fig. 2f)��������������������������������������������3 (Fig. 11a) ����������������������������������������������� fumida van der Vecht
3 Gena more than 1.7× medial width of compound eye in side - Clypeus not bulging medially, flat or gently curved in side view
view (Fig. 3a), interocellar distance much less than the distance (Fig. 11b)������������������������������������������������������������������������������15
from ocelli to the vertex���������������������������������������������������������4 15 Metasomal segments 3–5 mostly yellow (Fig. 11c); vertex
- Gena less than 1.7× medial width of compound eye (Fig. 3b), orange-yellow to red ���������������������������������������crabro Linnaeus
interocellar distance as long or almost as long as the distance - Metasomal segments 3–5 mostly dark brown or black, some
from ocelli to the vertex ��������������������������������������������������������5 segments with narrow distal yellow band (Fig. 11d); vertex red-
4 Metasomal terga 3–6 black, or at most with narrow apical dish to dark brown���������������������������������������� dybowskii André

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band on tergum 3 (Fig. 3c)���������������������������� soror du Buysson 16 Clypeus (Fig. 12a) and metanotum with dark markings
- Metasomal segments 3–6 with orange apical band, tergum 6 ���������������������������������������������������������������� multimaculata Pérez
mostly orange (as in Fig. 3d) ���������������������� mandarinia Smith - Clypeus and metanotum without black markings (Fig. 12b and
5 Punctures on sides of tergum 2 large, separated by less than one d)������������������������������������������������������������������������������������������17
puncture width (Fig. 4a); pronotal carina barely interrupted by 17 Clypeus yellow with black markings (Fig. 12a), or if without
pronotal fovea (Fig. 4c)����������������������������������������������������������6 markings then metanotum entirely yellow (as in Fig. 12c and d)
- Punctures on sides of metasomal tergum 2 small, separated by ���������������������������������������������������������������� bellicosa de Saussure
more than one puncture diameter (Fig. 4b); pronotal carina - Clypeus yellow to brown, without black markings (Fig. 12b);
widely interrupted by pronotal fovea (Fig. 4d)�����������������������7 metanotum coloration variable (Fig. 12d and e)�������������������18
6 Clypeus mostly black (Fig. 4e); metasomal terga mostly black 18 Scutellum and metanotum entirely or primarily yellow
or at most with yellow apical band on tergum 1 (Fig. 5a) (Fig. 12d) and vertex black�����������������������������bicolor Fabricius
����������������������������������������������������������������������������fervida Smith - Scutellum and metanotum not primarily yellow (Fig. 12e), or if
- Clypeus yellow (Fig. 4f); metasomal terga 1–5 often with apical primarily yellow then vertex only partly black����������������������19
yellow band (Fig. 5b) �������������������������������luctuosa de Saussure 19 Scutal punctures small, separated by 2–3 puncture diameters
7 Pretegular carina complete (Fig. 7a); clypeus medially with (Fig. 13a); metasomal terga 2–5 entirely black����� basalis Smith
coarse or large punctures, separated by one puncture diameter - Scutal punctures large, separated by 1 puncture diameter or less
or less (Fig. 6c)�����������������������������������������������������������������������8 (Fig. 13b); metasomal terga 2–5 not entirely black���������������20
- Pretegular carina incomplete (Fig. 7b); middle of clypeus with 20 Clypeal punctation uniformly distributed�������� simillima Smith
small punctures separated more than one puncture diameter - Clypeal punctation irregularly distributed, with impunctate
(Fig. 6d)��������������������������������������������������������������������������������16 areas�������������������������������������������������������������������������������������21
8 Apical margin of clypeus produced into two acute triangular 21 Side of prothorax close to spiracle smooth (Fig. 13c); vertex not
projections (Fig. 8a)����������������������������������������������������������������9 black; mesothorax and propodeum with same coloration; terga
- Apical margin of clypeus produced into two broadly rounded 2 and 5 black or mostly black������������������������������� vivax Smith
projections (Fig. 8b)�������������������������������������������������������������11 - Side of prothorax close to spiracle rugose (Fig. 13d); without
9 Vertex black (Fig. 8c) and first three metasomal terga primarily above color combination ………..………...…velutina Lepeletier
orange-yellow (Fig. 7a)��������������������philippinensis de Saussure
- Vertex orange (Fig. 8d) or light brown, if black then orange-
yellow coloration restricted to first two metasomal terga
(Fig. 7c), or first three metasomal terga black, with at most World Species of Vespa
narrow distal orange-yellow band (Fig. 7d)��������������������������10 Vespa affinis (Linnaeus)
10 Metasomal terga never entirely black; metasomal tergum 2 Figs. 4d, 9c, 10b, 10d, and 14
with broad orange-yellow band extending one-third or more
across tergum (Fig. 7c and d)�������������������������������ducalis Smith Apis affinis Linnaeus, 1764: 417. Holotype female; ‘in Calidis
- Metasomal terga all black (Fig. 7f), or metasomal tergum 2 regionibus’ (UPPSALA).
orange-yellow (Fig. 7e), or tergum 2 black with narrow, distal Vespa affinis Fabricius, 1787: 287. Syntype females; China
orange-yellow band that extends much less than one-third of (COPENHAGEN). Nec Vespa affinis (Linnaeus, 1764).
tergal width (Fig. 7b).………..........………tropica (Linnaeus) Vespa unifasciata Olivier, 1792: 677. Type unknown; ‘Indes
11 Metasomal terga 2, 5 and 6 reddish brown or dark brown; orientales’ (repository unknown). Nec Vespa unifasciata Gmelin,
tergum 3 and usually tergum 4 mostly yellow, with basal red- 1790.
dish brown band extending medially and two small, lateral red- Vespa alduini Guérin-Méneville, 1831: 264. Holotype male;
dish brown spots (Fig. 9a and b)���������������� orientalis Linnaeus Indonesia, Maluku, Buru Isl., ‘l’île de Bourou, l’une des Moluques’
- Metasomal terga coloration not as above�����������������������������12 (GENOA).
12 Metasomal segment 1 0.5× as long as or longer than width in Vespa bimaculata Guérin-Méneville, 1838: 264. Unnecessary re-
dorsal view (Fig. 9c)�������������������������������������������������������������13 placement name for Vespa alduini Guérin-Méneville. Nec Vespa
- Metasomal segment 1 less than 0.5× as long as wide in dorsal bimaculata Geoffroy, 1785, and Vespa bimaculata Olivier, 1792.
view (Fig. 9d)�����������������������������������������������������������������������14 Vespa nigripennis de Saussure, 1854: 156. Type unknown; Philippines
13 Scutellar punctures small and separated two puncture diam- (repository unknown). Nec Vespa nigripennis Degeer, 1773.
eters (Fig. 10a); metasomal tergum 2 mostly black (Fig. 10c) Vespa cincta var. picea du Buysson, 1905 (1904): 537, Lectotype
����������������������������������������������������������� mocsaryana du Buysson female (designated by van der Vecht, 1959: 214); Papua New
- Scutellar punctures large and contiguous or separated by 1 Guinea, ‘Tupuseleia or Kapakapa’ (van der Vecht does not indi-
puncture diameter or less (Fig. 10b); metasomal tergum 2 cate which locality) (GENOVA).
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Figs. 5–8. Vespa morphology. Fig. 5. Lateral view of metasomal tergum 2. Fig. 6. (a, b) Lateral view of pretegular pronotal carina. (c, d) Front view of clypeus. Fig.
7. (a, b) Front view of clypeus. (c, d). Dorsal view of vertex. Fig. 8. (a, c) Lateral view of metasoma. (b, d, e) Dorsal view of metasoma. Fig. 5a, V. fervida. Fig. 5b,
V. luctuosa. Figs. 6a, 6c, 7a, 8a, 8c, V. philippinensis. Figs. 6b, 6d, V. velutina. Figs. 7b, 7e, 7f V. tropica. Figs. 7c, 7d, 8d, V. ducalis. Fig. 8b, V. bicolor.

Vespa indosinensis Pérez, 1910: 8. Lectotype female (designated by Vespa affinis archboldi van der Vecht, 1957: 32. Holotype female;
van der Vecht, 1957: 28); Vietnam: Annam (PARIS). Given as V. Indonesia: ‘Hollandia’ (LEIDEN).
affinis indosinensis by van der Vecht, 1957. Vespa affinis moluccana van der Vecht, 1957: 32. Holotype female;
Vespa formosana Sonan, 1927: 125. Lectotype female (designated by Indonesia: Saparua I. (LEIDEN).
Kojima et al. 2011: 45); Taiwan: Taihoku (TAICHUNG). Vespa affinis alticincta van der Vecht, 1957: 33. Holotype female;
Vespa affinis var. continentalis Bequaert, 1936: 350. Holotype fe- New Britain (LONDON-NHM).
male; India: Mangalore (WASHINGTON).
Distribution. India, Sri Lanka, Bangladesh, China, Hong Kong,
Vespa affinis var. hainanensis Bequaert, 1936: 349. Holotype female;
Taiwan, Japan (Ryukyu Is.), Myanmar, Thailand, Laos, Vietnam,
China: Hainan Is., 2 mi s Nodoa (CAMBRIDGE).
Malaysia, Singapore, Indonesia, Papua New Guinea, Philippines,
Vespa affinis nigriventris van der Vecht, 1957: 29. Holotype female;
and introduced into Australia, New Zealand, United States.
Philippines: Palawan, Puerto Princesa (LOGAN).
Vespa affinis rufonigrans van der Vecht, 1957: 29. Holotype female; Discussion. This is one of the smaller bodied hornets, and is common
Indonesia: Sulawesi, Palu ‘North West Celebes’ (LEIDEN). in subtropical and tropical Asia. They are generalist scavengers,
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Figs. 9–12. Vespa morphology. (a, b) Dorsal view of metasoma. (c, d) Dorsal view of basal metasomal terga. Fig. 10. (a, b) Dorsal view of scutum. (c, d) Dorsal
view of metasoma. Fig. 11. (a, b) Lateral view of head. (c, d) Dorsal view of metasoma. Fig. 12. (a, b) Front view of clypeus. (c-f) Dorsal view of metanotum. Figs.
9a, 9b, V. orientalis. Figs. 9c, 10b, d, V. affinis. Fig. 9d, V. crabro. Figs. 10a, 10c, 11b, 11c, V. mocsaryana. Fig. 11a, V. fumida. Fig. 11d. V. dybowskii. Figs. 12a, 12f,
V. multimaculata. Figs. 12b, d, e, V. bicolor. Fig. 12c, V. luctuosa.
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Figs. 13 and 14. Vespa morphology. (a, b) Dorsal view of scutum. (c, d) Lateral view of pronotal lobe. (e, f) Lateral view of anterior pronotal carina. (a) V. basalis.
(b) V. ducalis. (c) V. vivax. (d) V. velutina. Fig. 14. V. affinis coloration. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

feeding on nectar, fruit, tree sap, carrion, and insects, including Vespa tridentata Cameron, 1903: 278. Lectotype female (designated
honeybees. Vespa affinis nests are usually built high in trees, but can by Kojima, 1997: 20); Japan (LONDON-NHM).
also be found in shrubs, and on and in buildings (Archer 1997). Vespa nigrans du Buysson, 1903: 175. Holotype male; China: ‘Yun-
nam, Tsé-kou’ (PARIS).
Vespa analis Fabricius Vespa parallela var. biroi du Buysson, 1905 (1904): 513. Lectotype
Figs. 2d, 2e, 15, and 16 female (designated by van der Vecht 1957); Singapore (PARIS).
Vespa analis Fabricius, 1775: 363. Holotype female; ‘in Cap. B. S’. Vespa analis var. tenebrosa du Buysson, 1905 (1904): 516. Lectotype
[in error, probably Java] (LONDON-LS). female (designated by van der Vecht, 1957: 14); Indonesia: Java,
Vespa crabro sphinx Christ, 1791: 217. Holotype female (destroyed). Goban (PARIS).
Vespa tyrannica Smith, 1857:119. Lectotype female (designated by Vespa analis var. (or subsp.) barbouri Bequaert, 1939: 40. Holotype
van der Vecht, 1959: 215); Singapore (LONDON-NHM). female; India: Sikkim, Teesta Valley (CAMBRIDGE).
Vespa japonica Smith, 1868: 279. Lectotype male (designated by Vespa analis var. (or subsp.) kuangsiana Bequaert, 1939: 42.
Kojima, 1997: 20); Japan: ‘Hakodadi’ (LONDON-NHM). Nec Holotype female; China: ‘Kwangsi’, (CAMBRIDGE).
Vespa japonica Radoszkowski, 1857, and Vespa japonica de Vespa analis eisa Yamane, 1987: 631. Holotype female; Japan:
Saussure, 1858 (=Vespula flaviceps (Smith)). ‘Yona, Okinawa-jima’ (KAGOSHIMA).
Vespa parallela André, 1884: lxi. Holotype female; Russia: Vespa analis nagatomii Yamane, 1987: 632. Holotype female; Japan:
Vladivostock, ‘sur l’Amour’ (PARIS). ‘Koza-dake, Iriomote-jima’ (KAGOSHIMA).
Vespa insularis Dalla Torre, 1894, Cat. Hym. 9: 147. Replacement Vespa maguanensis Dong, 2001: 82. Holotype female; China:
name for Vespa japonica Smith. Yunnan (KUNMING). Synonymized by Carpenter et al. (2011).
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Figs. 15 and 16. Vespa analis color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

Vespa hekouensis Dong and Wang, 2003: 407. Holotype female; Vespa basalis Smith
China, Yunnan, Hekou County (KUNMING). Synonymized by Figs. 13a and 17
Carpenter et al. (2011).
Vespa basalis Smith, 1852: 46. Holotype female; Nepal: ‘Nepaul’
Distribution. India, Nepal, China, Hong Kong, Taiwan, Korea, (LONDON-NHM).
Japan, Russia, Myanmar, Thailand, Laos, Vietnam, Malaysia, Vespa obliterata Smith, 1852: 47. Holotype female; ‘Northern India’
Malaya, Singapore, Indonesia. (LONDON-NHM).
Vespa basilis Dover, 1929: 48. Misspelling of Vespa basalis Smith,
Discussion. This is one of the most widely distributed species of 1852.
Vespa. Its native range includes tropical Asia and extends north into
Japan, Russia, and Korea. Nests are typically built in trees 2–3 m Distribution. Pakistan; India, Nepal, China, Taiwan, Myanmar,
above the ground (Archer 1998b). Thailand, Laos, Vietnam, Indonesia: Sumatra.
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Figs. 17 and 18. Vespa color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face. Fig. 17. V. basalis. Fig. 18. V. bellicosa.

Discussion. Vespa basalis occurs in forested regions between 500 m black and yellow banding on the thorax and metasoma. It is not
and 1,500 m (Archer 1999). Nests are built on tree branches, in known where V. bellicosa build their nests.
shrubs, on buildings, in rock crevices, and even in the ground. The
mostly dark brown to black metasoma is one of the more distinctive Vespa bicolor Fabricius
features of V. basalis. Figs. 2f, 8b, 12b, 12d, 12e, 19, and 20

Vespa bellicosa de Saussure Vespa bicolor Fabricius, 1787: 288. Syntypes, sex not stated; China
Fig. 18 (COPENHAGEN).
Vespa lutea Coquebert, 1804: 94. Type unknown; ‘Massiliae lecta, in
Vespa bellicosa de Saussure, 1854: 146. Lectotype female (desig-
navi ex India’ (repository unknown).
nated by van der Vecht, 1959: 216); ‘Java’ [error; Sumatra or
Vespa auraria var. citriventris du Buysson, 1905 (1904): 552.
Borneo] (TURIN).
Lectotype female (designated by van der Vecht, 1959: 218);
Distribution. Indonesia: Sumatra, Borneo (Kalimantan, Sarawak). Sikkim: Padamchen Lingtou, ‘Padamtsin à Lingtou’ (PARIS).
Discussion. This species appears to be associated with lowland trop-
ical forests in Borneo and Sumatra (Archer 1999). It is unlikely that Distribution. India, Bhutan, Nepal, China, Hong Kong, Myanmar,
it would become invasive in other regions. It has very distinctive Thailand, Laos, Cambodia, Vietnam. Introduced into Taiwan.
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Figs. 19 and 20. Vespa bicolor color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

Discussion. This species of Vespa has a very distinctively marked Distribution. India, Myanmar, Thailand, Laos, China, Russia, Korea.
black and yellow thorax, with a largely yellow metasoma. It preys Discussion. This is a high altitude species, found between 200 m
on honeybees, and one species of Dendrobium orchid exploits this and 2,000 m (Archer 1999). Zhang (1989) reported the presence of
behavior. These orchids produce chemicals that mimic alarm phero- Vespa binghami in Miocene deposits in Shandong, China, but this
mones of Asian (Apis cerana) and European (Apis mellifera) honeybees identification needs to be verified.
to attract the hornets to pollinate their flowers (Brodman et al. 2009).

Vespa crabro Linnaeus

Vespa binghami du Buysson
Figs. 1, 2a, 2b, 3b, 3d, 9d, 11b, 11c, and 22–26
Figs. 2c and 21

Vespa binghami du Buysson, 1905 (1904): 523. Lectotype female Vespa crabro Linnaeus, 1758: 572. Holotype female; ‘in Europae’
(designated by van der Vecht, 1959: 210); Myanmar: Toungoo, (LONDON-LS).
‘Taungoo Hills’ (PARIS). Vespa vexator Harris, 1776: 128. Holotype female; ‘English’
Vespa suprunenkoi Birula, 1925 (1924): 92. Lectotype female (destroyed).
(designated by Kurzenko, 2004: 197); Korea: Sakhalin Is. (ST. Vespa crabro major Retzius (in Degeer), 1783: 63; Type unknown
PETERSBURG). (repository unknown).
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Figs. 21 and 22. Vespa color forms, (a) Lateral view. (b) Dorsal view. (c) Front view of face. Fig. 21. Vespa binghami. Fig. 22. Vespa crabro.

Vespa pratensis Geoffroy (in Fourcroy), 1785: 437; Type unknown; Vespa oberthuri du Buysson, 1902: 140; Syntype females; China:
France: ‘in Agro Parisiensi’ (repository unknown). ‘Chine: Se-Tchouen, Sio-Lou’ (PARIS).
Vespa crabro germana Christ, 1791: 215. Type unknown (destroyed). Vespa flavofasciata Cameron, 1903: 280. Lectotype female (des-
Vespa crabroniformis Smith, 1852: 40. Syntype female, male; ‘North ignated by Kojima, 1997: 21); Japan: ‘Nügata (Shinanogawa)’
China’ (LONDON-NHM). (LONDON-NHM).
Vespa crabro var. borealis Radoszkowski, 1863: 128. Syntype fe- Vespa crabro var. tartarea du Buysson, 1905 (1904): 506. Syntype
male, male; Russia: ‘Pargolova i Osinovoa Roshchi’ (reposi- females; Japan: ‘Yokohama ... Columbia’ (BUDAPEST).
tory unknown). Nec Vespa borealis Kirby, 1837, Vespa borealis Vespa crabro var. altaica Pérez, 1910: 5. Holotype Female; Russia:
Zetterstedt, 1840, and Vespa borealis Smith, 1843. Altaï (PARIS).
Vespa crabro var. anglica Gribodo, 1892 (1891): 242. Syntype females; Vespa crabro var. caspica Pérez, 1910: 6. Holotype Female;
England: ‘Inghilterra’ (GENOA). Nec Vespa anglica Smith, 1843. Azerbaijan: ‘Talysch et Lenkoran, région Caspienne’ (PARIS).
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Figs. 23 and 24. Vespa crabro color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

Vespa crabro nigra Birula, 1925: 55. Syntypes female, male; Russia: Vespa crabro var. birulai Bequaert, 1931: 105. Replacement name
West Siberia from Urals to Enisei and slightly after Enisei [in for Vespa crabro chinensis Birula.
Russian] (ST. PETERSBURG). Nec Vespa nigra Geoffroy, 1785. Vespa crabro var. gribodoi Bequaert, 1931: 105. Replacement name
Vespa crabro vulgata Birula, 1925: 55. Syntype female, male: west for Vespa crabro var. anglica Gribodo.
Europe (St. Petersburg).
Vespa crabro meridionalis Birula, 1925: 55. Syntype females: Distribution. Eurasia, Algeria. Introduced into eastern North
Transcaucasia, north Persia, western part of Transcaspian region America, and Guatemala.
(ST. PETERSBURG).
Vespa crabro chinensis Birula, 1925: 55. Syntypes female, male: Discussion. Vespa crabro usually nests in sheltered aboveground
middle and south China [in Russian] (ST. PETERSBURG). Nec sites, such as tree hollows, wall voids, beehives, and outhouses
Vespa chinensis Fabricius, 1793. (Archer 1993). However, nests have been found in subterranean
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Figs. 25 and 26. Vespa crabro color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

sites as well. This species adapts well to urban and suburban set- Vespa ducalis Smith, 1852: 39. Syntypes female, male; China: ‘Tein-
tings. It is widespread in Eurasia, and now eastern North America. tung, near Nigo-po-foo’ (LONDON-NHM).
The species was first reported in the United States in New York in Vespa ducalis var. pulchra du Buysson, 1905 (1904): 519. Lectotype
the 1800s. More recently, the species was recorded from a single female (designated by van der Vecht, 1959: 224); Japan:
worker collected in Guatemala City, Guatemala. It was found in the Yokohama (PARIS).
Entomological Collection of the Universidad del Valle de Guatemala Vespa matsumurai Sonan, 1935: 370. Holotype female; Japan:
(Landolt et al. 2010). There is no evidence that the species is estab- Tokyo (TAICHUNG).
lished in Guatemala. Nests are built in cavities in trees or buildings. Vespa esakii Sonan, 1935: 371. Holotype female; Japan: Tsushima
Is., ‘Izuhara’ (FUKUOKA).
Vespa ducalis Smith Vespa tropica loochooensis Bequaert, 1936: 343. Holotype female;
Figs. 7c, 7d, 8d, 13b, 27, and 28 Japan: Iriomote Is., Ryukyu Archipelago (CAMBRIDGE).
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Figs. 27 and 28. Vespa ducalis color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

Vespa tropica pseudosoror van der Vecht, 1959: 224. Holotype Vespa dybowskii ‘Rad. in litt’. André, 1884: 582. Holotype female;
female; Vietnam: Annam, Tourane, 1,000 m (PARIS). Russia: ‘Sibérie’ (PARIS?).
Vespa dubowskii; Dalla Torre, 1904: 65. Misspelling of Vespa
Distribution. India, Sikkim, Nepal, Myanmar, Thailand; Laos,
dybowskii André.
Vietnam, China, Hong Kong, Hainan; Taiwan; Russia: Korea; Japan,
Vespa walkeri du Buysson, 1905 (1904): 539. Syntype female, male;
including Ryukyu Is.
China: ‘Haï-ning’ (LONDON-NHM).
Discussion. This species of Vespa is quite different from the others. It Vespa dybowskii mutata Ma, 1937: 30. Holotype male; China:
preys on nests of other paper wasps, feeding the paper wasp pupae ‘Hangchow’ (HANGZHOU, destroyed?).
and larvae to their larvae. Their colonies are small with an average Distribution. Myanmar, China (Tibet, Zhejiang), Russia (Siberia,
of 50 individuals, and nests are built underground, in tree hollows, Primorsky), Korea, Japan.
and even in attics (Archer 1991). Discussion. This species is relatively rare in collections. It is a nest
parasite of Vespa crabro and V. simillima. Vespa dybowskii queens
Vespa dybowskii André take over the nests of these species and use the host workers to
Figs. 11b, 11d, and 29 raise their own brood (Martin et al. 2008)
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Figs. 29 and 30. Vespa color forms, (a) Lateral view. (b) Dorsal view. (c) Front view of face. Fig. 29. V. dybowskii. Fig. 30. V. fervida.

Vespa fervida Smith Vespa variabilis fumida van der Vecht, 1959: 228. Holotype female;
Figs. 4a, 4e, 5a, and 30 India: ‘British Bootan’, Padong’ (PARIS).

Vespa fervida Smith, 1858: 23. Holotype female; Indonesia: Sulawesi, Distribution. India, Bhutan, Nepal, Myanmar, China (Sichuan,
‘Celebes’ (LONDON-NHM). Yunnan, Hubei, Fujian).
Discussion. Vespa fumida is found at high altitudes between 600 m
Distribution. Indonesia (Sulawesi, Salayar, Buton). and 3,000 m across northeastern India into China (Archer 1999).
Discussion. This species has a very limited distribution on Sulawesi Otherwise, little is known about its biology.
and nearby islands in Indonesia. They build nests in forested areas,
and around human habitations, nests have been found under leaves
Vespa luctuosa de Saussure
and on fences. It is a small-bodied, dark-colored Vespa. Overall, its
Figs. 4c, 4f, 5b, 12c, and 32
biology is poorly known (Matsuura and Yamane 1984, Archer 1999)
Vespa luctuosa de Saussure, 1854: 143. Holotype male; Philippines
Vespa fumida van der Vecht (LONDON-NHM).
Fig. 31 Vespa bellicosa var. semperi du Buysson, 1905 (1904): 542. Lectotype
female (designated by van der Vecht, 1957: 48). Philippines:
Vespa variabilis du Buysson, 1905 (1904): 522. Lectotype female Camiguin Is., ‘Comigu’ (VIENNA).
(designated by van der Vecht, 1959: 228); China: Mou-pin Vespa luctuosa var. luzonensis Bequaert, 1934: 5. Holotype female;
(PARIS). Nec Vespa variabilis Fabricius, 1781. Philippines: Luzon (BERLIN).
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Figs. 31 and 32. Vespa color forms, (a) Lateral view. (b) Dorsal view. (c) Front view of face. Fig. 31. Vespa fumida. Fig. 32. V. luctuosa.

Vespa luctuosa negrosensis Kojima and Tano, 1985: 436. Holotype Vespa japonica Radoszkowski (in Motschulsky), 1857: 410. Type?;
female; Philippines: ‘Mambucal, Negros Is’. (TOKYO). Japan, ‘Japon’ (ST. PETERSBURG).
Synonymized by Archer (1999). Vespa bellona Smith, 1871: 248.Holotype female; China: ‘Yunan’
(LONDON-NHM).
Distribution. Philippines.
Vespa magnifica var. latilineata Cameron, 1903: 278. Holotype fe-
Discussion. This species has been found nesting in forested regions be- male; Japan: ‘Hitoyoshi’ (LONDON-NHM).
tween 200 and 1,500 m elevation. The nests are built on tree branches, Vespa mandarina; Dalla Torre, 1894: 149. Misspelling of Vespa
sometimes as high as 20 m above the ground (Archer 1999). mandarinia Smith.
Vespa magnifica var. nobilis Sonan, 1929: 140. Holotype female;
Taiwan: ‘Musha’ (TAICHUNG).
Vespa mandarinia Smith
Vespa magnifica sonani Matsumura, 1930: 1. Lectotype female (des-
Figs. 33–35 ignated by Kojima, 1997: 22); Taiwan: ‘Sina’ (SAPPORO).
Vespa mandarinia Smith, 1852: 38. Holotype female; China: ‘Tein-
tung, near Ning-po-foo’ (LONDON-NHM). Distribution. India, Sri Lanka, Bhutan, Nepal, Myanmar, Thailand,
Vespa magnifica Smith, 1852: 45. Syntype females; Nepal ‘Nepaul’ Laos, Vietnam, Malaysia, Malaya, China, Hong Kong, Taiwan,
(LONDON-NHM, OXFORD). eastern Russia, Korea, Japan (including Ryukyus).
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Figs. 33–35. Vespa mandarina color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

Discussion. This species occurs in hilly regions. It nests in the ground, Vespa multimaculata Pérez, 1910: 14. Lectotype female (designated
starting with mammal burrows or decayed root cavities, enlarging by van der Vecht 1957); Brunei (PARIS).
them as the colony develops (Archer 1995, 2008). This is another Vespa luctuosa var. malayana Bequaert, 1934: 4. Replacement name
species of Vespa that preys on honeybees. Zhang (1989) reported for Vespa annulata Smith.
the presence of Vespa magnifica in Miocene deposits in Shandong, Vespa multimaculata pendleburyi van der Vecht, 1957: 43; Holotype
China. This identification needs to be verified. This species has also female; Malaysia: ‘near Jitra, Kedah’, (LEIDEN).
been introduced into the Pacific Northwest, where nests have been
found recently (Anonymous 2020). Distribution. Thailand; Laos; Malaysia: Peninsular Malaysia,
Sarawak, Sabah; Singapore; Indonesia: Sumatra, Borneo
Vespa mocsaryana du Buysson (KALIMANTAN); Brunei.
Figs. 10a, 10c, 36, and 37
Discussion. Vespa multimaculata is found in lowland and montane
Vespa mocsaryana du Buysson, 1905 (1904): 537. Syntype females,
forests in Southeast Asia (Archer 1999, 2011), and is unlikely to dis-
males; India: Malaysia: ‘Presqu’ile de Malacca: Pérak; India:
perse to temperate regions. Martin (1995) found nests in the ground
Sikkim; Assam: Khasia Hills; Myanmar: Tenasserim: Thagatâ’
(BUDAPEST, VIENNA, LONDON-NHM, GENOVA). beneath the roots of a tree.

Distribution. India: Meghalaya, Sikkim, Assam; China: Sichuan,

Vespa orientalis Linnaeus
Anhui, Fujian, Hong Kong; Myanmar; Thailand; Laos; Vietnam;
Figs. 9a, 9b, and 40
Malaysia: Peninsular Malaysia; Indonesia: Sumatra.
Vespa orientalis Linnaeus, 1771: 540. Holotype female; ‘Oriente’
Discussion. Not much is known about the biology of this species. It
(LONDON-LS).
occurs in montane forests in southern Asia. Nests have been found
Vespa turcica Drury, 1773: 74. Type unknown; Turkey: Smyrna
in a shrub and on a ceiling (Archer 2008).
(destroyed?).
Vespa multimaculata Pérez Vespa quadripunctata Forskål, 1775: 84; Type?; Egypt: Cairo,
‘Kahirae’ (COPENHAGEN?).
Figs. 4b, 12a, 12f, 38, and 39
Vespa crabro fusca Christ, 1791: 216. Type unknown; Turkey:
Vespa annulata Smith, 1858: 116. Lectotype female (designated ‘Smirna’ (destroyed).
by Carpenter and Kojima 1997); Malaysia: Borneo: Sarawak Vespa aegyptiaca Vallot, 1802: 170; Type unknown; Egypt (reposi-
(OXFORD). Nec Vespa annulata Rossi, 1790. tory unknown).
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Figs. 36 and 37. Vespa mocsaryana color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

Vespa nilotica Vallot, 1802: 170; Type unknown; Egypt (repository Discussion. This is a distinctively marked reddish orange and yellow
unknown). wasp. The nests are built in the ground in rodent burrows or rock
Vespa jurinei de Saussure, 1854: 133; Lectotype female (designated crevices (Archer 1998a). Around human habitation, nests have also
by Guiglia, 1971: 94); Albania (LONDON-NHM). been found in empty beehives, under eaves or floorboards, and even
Vespa orientalis var. aegyptiaca André, 1884: 584’ Type unknown; in abandoned outhouses. It has been recorded from Mexico but is
Egypt: Cairo, Alexandria ‘Le Caire, Alexandrie’ (PARIS). Nec evidently not established (Dvořák 2006). Vespa orientalis are scaven-
Vespa aegyptiaca Vallot, 1802. gers, feeding on nectar and fruit as well as insects and carrion. They
Vespa indica Wroughton, 1889: 35. Lapsus for Vespa orientalis also prey on honeybees and other pollinators. A study by Plotkin
Linnaeus. et al. (2010) found that this species has a novel form of photosyn-
Vespa orientalis var. zavattarii Guiglia and Capra, 1933: 168. Syntype thesis using yellow pigments in the cuticle.
females; Libya: ‘Fezzan, Ubari; Algeria: Oued Tizzi’ (GENOA).
Vespa orientalis var. somalica Giordani Soika, 1934: 184. Lectotype
Vespa philippinensis de Saussure
female (designated by Giordani Soika, 1973: 11); Somalia:
Figs. 6a, 6c, 7a, 8a, 8c, and 41
‘Somalia italiana di Carim’ (VENICE).
Vespa orientalis arabica Giordani Soika, 1957: 482. Holotype Vespa philippinensis de Saussure, 1854: 148. Holotype male;
female; Yemen: ‘Western Aden Protectorate: Al Milah’ Philippines (LONDON-NHM).
(LONDON-NHM).
Distribution. Philippines (Leyte, Luzon, Negros, Samar).
Distribution. north Africa, southern Europe, Turkey, Middle
East, southern Russia, Afghanistan, Pakistan, India, Nepal, China Discussion. Little is known about the biology of this endemic spe-
(Xinjiang). Introduced into Czech Republic; Spain; Madagascar; cies. A nest described by Starr (1987) from the island of Leyte was
Mexico. found in a cavity in the ground (Archer 1991).
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Figs. 38 and 39. Vespa multimaculata color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

Vespa simillima Smith Discussion. Nests of this species have been found in the ground, and
Figs. 42 and 43 in sheltered above ground sites, such as hollow trees, wall voids, and
attics. A queen of V. simillima was found in the village of Tsaochan
Vespa simillima Smith, 1868: 280. Holotype female; Japan: in Taiwan in 2003, but it is unclear whether the species has estab-
‘Hakodadi’ (LONDON-NHM). lished on the island (Sung et al. 2006). It is apparently not estab-
Vespa mongolica André, 1884: 4. Syntype male, female; Russia: lished in British Colombia (Kimsey and Carpenter 2012).
‘Vladivostock, sur l’Amour, dans la Sibérie orientale’ (PARIS?).
Vespa xanthoptera Cameron, 1903: 278. Holotype male; Japan:
Vespa soror du Buysson
‘Michzusawa’ (LONDON-NHM).
Figs. 3a, 3c, and 44
Vespa micado Cameron, 1903: 279. Lectotype female (designated by
Kojima, 1997: 20); Japan: Nagasaki (LONDON-NHM). Vespa ducalis var. soror du Buysson, 1905 (1904): 519. Lectotype female
Vespa mongolica var. sexpunctata Pérez, 1905: 24. Holotype female; (designated by van der Vecht, 1957: 16); China: Jaingxi, ‘Kiang-si’
Japan: ‘Yokohama’ (PARIS). (PARIS). Given as mandarina soror by van der Vecht, 1957.
Vespa mongolica var. flavata Pérez, 1910: 17. Holotype female;
China (PARIS). Distribution. India, southwestern China, Hong Kong, Thailand,
Laos, Vietnam.
Distribution. China (Liaoning), southwestern Russia, Korea, Japan
(including Yaku-shima Is.). Introduced into British Columbia, Discussion. Little biology is known for this species. It has been found
Canada, but not established; Taiwan? in hilly and montane sites up to 1,500 m in continental southeastern
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Figs. 40 and 41. Vespa color forms, (a) Lateral view. (b) Dorsal view. (c) Front view of face. Fig. 40. V. orientalis. Fig. 41. V. philippinensis.

Asia (Archer 2008). The nests are typically underground. These Vespa unicolor Smith, 1863: 44. Holotype female; Indonesia: ‘Bouru’
wasps are predators of other insects and even small vertebrates, like (OXFORD).
geckos. They will also attack nests of honeybees and other social Vespa cinta Wroughton, 1889: 35. Misspelling of Vespa cincta
vespids, including hornets (Lee 2009). Degeer 1773.
Vespa eulemoides du Buysson, 1905 (1904): 530. Holotype female;
Vespa tropica (Linnaeus) India: ‘Ile Andaman: port de Blair’ (GENOA).
Figs. 7b, 7e, 7f, and 45–49 Vespa affinis var. trisignata Pérez, 1910: 8. Holotype female;
Indonesia: ‘Timor’ (PARIS).
Sphex tropica Linnaeus, 1758: 571. Holotype female; ‘in Indiis’ Vespa rubricans Pérez, 1910: 10. Lectotype female (designated
(UPPSALA). by van der Vecht 1957); ‘Lindi (Afrique orientale allemande)’
Vespa cincta Fabricius, 1775: 362. Holotype?; India: ‘ad littora (PARIS).
Malabarica’ (LONDON-LS). Nec Vespa cincta Drury, 1773. Vespa tropica var. anthracina Bequaert, 1936: 341. Holotype female;
Vespa crabro tenebrionis Christ, 1791: 216. Type? (destroyed). Philippines: ‘Sibuwan’ (WASHINGTON).
Vespa deusta Lepeletier, 1836: 506. Lectotype female (designated by Vespa tropica var. haematodes Bequaert, 1936: 338. Holotype fe-
van der Vecht, 1959: 226); ‘Patrie inconnue’ (TURIN). male; India: ‘Kooloo’ (CAMBRIDGE).
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Figs. 42 and 43. Vespa simillima color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

Vespa tropica leefmansi van der Vecht, 1957: 19. Holotype female; Vespa velutina Lepeletier
Indonesia: ‘Solok, Padang, Sumatra’ (LEIDEN) Figs. 6b, 6d, 13d, and 50–52
Vespa tropica trimeres van der Vecht, 1957: 19; Holotype female;
Indonesia: Sulawesi, ‘Palu, West Celebes’ (LEIDEN). Vespa velutina ‘De Haan’ Lepeletier, 1836: 507. Holotype female;
Vespa tropica cebuana Kojima and Reyes, 1984: 260. Holotype fe- Indonesia: ‘Java’ (TURIN?).
male; Philippines: ‘Cebu City’ (TOKYO). Vespa auraria Smith, 1852: 46. Syntype females; ‘Northern India’
(LONDON-NHM, OXFORD).
Distribution. Afghanistan, Pakistan, India, Sri Lanka, Bhutan, Vespa crabro var. immaculata Morawitz, 1889: 161. Holotype fe-
Nepal, southeastern China, Hong Kong, Myanmar, Thailand, Laos, male; China: ‘Kansu. Nanpin’. (ST. PETERSBURG). Nec Vespa
Cambodia, Vietnam, Malaysia, Malaya, Borneo, Indonesia, Papua immaculata Gmelin, 1790.
New Guinea (including New Britain), Philippines; exotic in Guam. Vespa fruhstorferi Stadelmann, 1894: 89. Holotype female;
Discussion. This is a distinctively colored Vespa, with a black Indonesia: West-Java, Gunung Gede, ‘in einer Höhe, 8000 ft’.
metasoma, except for a broad yellow to orange stripe across most (Berlin?).
of the second segment (as in V. affinis), although some individuals Vespa velutina var. ardens du Buysson, 1905 (1904): 550. Syntype fe-
may lack this stripe. Vespa tropica are nest predators of other so- male, male; Indonesia: Lombok, Sapit, Sumbawa Islands. (Vienna,
cial wasps as well as other large-bodied insects, including honeybees Budapest).
(Archer 1991). Nests are either above ground in tree cavities or in Vespa auraria var. nigrithorax du Buysson, 1905 (1904): 553.
cavities in the ground. Around human habitations, they will build Lectotype female (designated by van der Vecht, 1957: 37), India:
nests under eaves, in shed, and attics. Darjiling (PARIS).
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Figs. 44 and 45. Vespa color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face. Fig. 44. V. soror. Fig. 45. V. tropica.

Vespa velutina var. celebensis Pérez, 1910: 12. Holotype female; Vespa velutina floresiana van der Vecht, 1957: 40. Holotype female;
Indonesia: Sulawesi, ‘Bua-Kraëng’ (PARIS). Indonesia: Flores Is., Labuan Badjo (LEIDEN).
Vespa velutina var. megei Pérez, 1910: 13. Holotype female; China: Vespa velutina pruthii van der Vecht, 1959: 228. Holotype female;
‘Koueï-Tchéou (Chine centrale)’ (PARIS). Synonymized by van India: Kashmir, Srinagar (CALCUTTA).
der Vecht, 1957: 37.
Vespa mongolica var. divergens Pérez, 1910: 16. Holotype female; Distribution. Afghanistan, Pakistan, India, Bhutan, Nepal, southern
Malaysia: Perak (PARIS). China, Hong Kong, Taiwan, Myanmar, Thailand, Laos, Vietnam,
Vespa flavitarsus Sonan, 1929: 142. Holotype female; Taiwan: Malaysia, Malaya, Indonesia. Introduced into Korea, Japan
‘Taiko-san (Kagi)’ (coll. M. Kato). (Tsushima I.), France, Belgium, United Kingdom, Spain, Portugal,
Vespa auraria flavitarsis Ma, 1937: 31. Misspelling of Vespa Italy, Yemen.
flavitarsus Sonan, 1929?
Vespa velutina variana van der Vecht, 1957: 37. Holotype female: Discussion. Vespa velutina has distinctive yellow tarsi and vel-
Thailand: Ban Umphang, Doi Hua Mot (LEIDEN). vety appearing dark brown to black thorax and metasoma. It is
Vespa velutina karnyi van der Vecht, 1957: 38. Holotype female; native across Asia. However, V. velutina was introduced into
Indonesia: Sumatra: Bandar Buru, Deli (LEIDEN). Europe apparently with Chinese pottery in the past couple of dec-
Vespa velutina sumbana van der Vecht, 1957: 40. Holotype female; ades (Keeling et al. 2017). Although this species preys on a var-
Indonesia: Sumba Is., Kananggar (LEIDEN). iety of insects, including flies, dragonflies, and Orthoptera, it is a
Vespa velutina timorensis van der Vecht, 1957: 40. Holotype female; notorious predator of pollinators, particularly honeybees (Apis
Indonesia: Timor Is., Mollo (BASEL). mellifera and A. cerana). Apis cerana have adapted to this hornet
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Figs. 46 and 47. Vespa tropica color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.

using avoidance behaviors and balling behaviors where they heat Conclusion
the hornet to death (Abrol 2006). From 2010 to 2018, there have been close to 50 interceptions of
Vespa (hornets) and Vespula (yellow jackets (Vespula) at US ports of
Vespa vivax Smith entry. Little less than half of those interceptions were hornets. The
Figs. 13d and 53 Vespa species intercepted include V. bellicosa, V. crabro, V. orientalis,
V. mandarinia, and V. tropica. One of the interceptions of signifi-
Vespa vivax Smith (in Horne and Smith), 1870: 190. Holotype fe- cance was an entire nest of V. mandarinia containing live brood and
male; India: Binsar, ‘Binsur, Kumaon’ (LONDON-NHM). pupae that was sent via express courier from Asia.
Vespa velutina var. mediozonalis Pérez, 1910: 14. Holotype female; All species of Vespa, except V. crabro, which is already intro-
India: Sikkim (PARIS). duced into the eastern United States, are considered of quarantine
Vespa wilemani Meade-Waldo, 1911: 104. Holotype female; Taiwan: importance by the USDA-APHIS. As part of the work of moni-
‘Formosa’ (LONDON-NHM). toring for possible introductions of hornets in the United States,
one of us (AHSP) created a website, Invasive Hornets (https://www.
Distribution. India, Nepal, Myanmar, Thailand, China (Sichuan,
ipmimages.org/browse/projectthumb.cfm?proj=1159), which is part
Yunnan, Tibet), Taiwan.
of a cooperation between the USDA, Animal Plant Health Inspection
Discussion. Little is known about this south Asian Vespa species. In
Service (APHIS), Plant Protection and Quarantine (PPQ) and the
Taiwan, nests have been found in montane regions at elevations be-
University of Georgia. This website contains more than 1,000
tween 1,500 and 2,500 m (Archer 1994).
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Figs. 48 and 49. Vespa tropica color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.
Insect Systematics and Diversity, 2020, Vol. 4, No. 3 25

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Figs. 50 and 51. Vespa velutina color forms. (a) Lateral view. (b) Dorsal view. (c) Front view of face.
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Figs. 52 and 53. Vespa color forms, (a) Lateral view. (b) Dorsal view. (c) Front view of face. Fig. 53. V. velutina. Fig. 54. V. vivax.

stacked, high-quality images of all the species and most of the races ages in this work and to PPQ- Science and Technology for supporting his
of the genus Vespa. work with bees and hornets. This is a contribution of the United States
It is important to have the resources for the identification and Department of Agriculture, Animal Plant and Health Inspection Service,
Plant Protection and Quarantine- Science and Technology, Identification
prevention of introduction of non-native species and to understand
Technology Program (ITP).
the potential effects of invasive hornets in our ecosystems. Hornets
are dangerous for the beekeeping industry because they can alter
pollination in agriculture and disrupt the beekeeping industry, as References Cited
well as create public health and safety problems. Anonymous. 2016. The greater banded hornet has found its way to Guam.
University of Guam Research and Extension. Available from https://
cnas-re.uog.edu/the-greater-banded-hornet-has-found-its-way-to-guam/.
Acknowledgments Anonymous. 2020. Three Asian giant hornets found in Nanaimo. Available from
We want to thank Christine Lebeau and Steve Heydon for helping to pro- https://bcinvasives.ca/news-events/news-from-iscbc/bc-government-
cess the loan of Vespa material from the AMNH and the BME, respect- bulletin-three-asian-giant-hornets-found-in-nanaimo
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