The recently rediscovered female holotype of Xylocopa coronata Smith, 1860, (Hymenoptera: Apidae) collected by Alfred Russel Wallace at the type locality of “Kaisaa” (= modern-day Kaioa, Maluku, Indonesia) is illustrated for the first time. Dorsal and lateral images of males and females of X. coronata and a map showing the known distribution of the species are provided. Females of this species exhibit geographic color variation in dorsal pubescence patterns and wing iridescence. At the present time, two subspecies are recognized: X. coronata combinata Ritsema from the island of Obi, Maluku, Indonesia, and X. coronata coro­nata Smith from additional islands (Bacan, Halmahera, Kaioa, Obi, Ternate, and Tidore) in the Maluku archipelago, Indonesia

Mawdsley, Jonathan R.

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9-25-2020 
Notes on the taxonomy and distribution of Xylocopa coronata 
Smith, 1860, a large carpenter bee from Maluku, Indonesia 
(Insecta: Hymenoptera: Apidae) 
Jonathan R. Mawdsley 
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Date of issue: September 25, 2020
0799
Notes on the taxonomy and distribution  
of Xylocopa coronata Smith, 1860,  
a large carpenter bee from Maluku, Indonesia  
(Insecta: Hymenoptera: Apidae)
Jonathan R. Mawdsley
Department of Entomology, MRC 187
National Museum of Natural History
Smithsonian Institution
10th Street and Constitution Avenue NW
Washington, DC, 20560 USA
Insecta MundIA journal of world insect systematics
Page Count: 5
Mawdsley JR. 2020. Notes on the taxonomy and distribution of Xylocopa coronata Smith, 1860, a large carpenter 
bee from Maluku, Indonesia (Insecta: Hymenoptera: Apidae). Insecta Mundi 0799: 1–5.
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Insecta MundI0799: 1–5 2020
Notes on the taxonomy and distribution  
of Xylocopa coronata Smith, 1860,  
a large carpenter bee from Maluku, Indonesia  
(Insecta: Hymenoptera: Apidae)
Jonathan R. Mawdsley
Department of Entomology, MRC 187
National Museum of Natural History
Smithsonian Institution
10th Street and Constitution Avenue NW
Washington, DC, 20560 USA
mawdsleyj@si.edu
Abstract. The recently rediscovered female holotype of Xylocopa coronata Smith, 1860, (Hymenoptera: 
Apidae) collected by Alfred Russel Wallace at the type locality of “Kaisaa” (= modern-day Kaioa, Maluku, 
Indonesia) is illustrated for the first time. Dorsal and lateral images of males and females of X. coronata and 
a map showing the known distribution of the species are provided. Females of this species exhibit geographic 
color variation in dorsal pubescence patterns and wing iridescence. At the present time, two subspecies are 
recognized: X. coronata combinata Ritsema from the island of Obi, Maluku, Indonesia, and X. coronata coro-
nata Smith from additional islands (Bacan, Halmahera, Kaioa, Obi, Ternate, and Tidore) in the Maluku 
archipelago, Indonesia. 
Key words. Geographic color variation, subspecies.
ZooBank registration. urn:lsid:zoobank.org:pub:0F05A789-DBAC-4944-A152-3C7FF2545606
Introduction
Xylocopa coronata Smith, 1860, is a brightly colored species of large carpenter bee (Hymenoptera: Apidae; Fig. 
1–12) which is generally distributed throughout the northern islands of the Maluku Archipelago of Indonesia 
(Fig. 13). The taxonomy of X. coronata was reviewed most recently by Lieftinck (1956), who provided keys to 
separate males and females of this species from those of sympatric species of Xylocopa but neither mapped the 
distribution of the species nor illustrated adult specimens. This paper provides new distribution records, maps, 
and color illustrations of males and females of this species, including the known geographic color forms of the 
females. I also report the rediscovery of the holotype specimen of X. coronata, which was missing from the Fred-
erick Smith collection at the time Lieftinck (1956) studied the species, because it was sent to Paul D. Hurd, Jr. of 
the Smithsonian Institution for his studies of Xylocopa (Hurd and Moure 1963).
Materials and Methods
The following notes are based on specimens of X. coronata contained in the entomological collections of the 
National Museum of Natural History, Smithsonian Institution (USNM). The late Smithsonian entomologist Paul 
D. Hurd, Jr. assembled a very large collection of pinned large carpenter bees from all areas of the world in order 
to support the preparation of his own monographic revisionary studies of Xylocopa (e.g., Hurd and Moure 1963; 
Hurd 1978). Specimens of X. coronata contained in this collection were authoritatively identified by previous 
workers, including Frederick Smith, Theodore D. A. Cockerell, Paul D. Hurd, Jr., and Maurits A. Lieftinck.  
2 · September 25, 2020 Mawdsley
Results
Xylocopa coronata Smith 1860: 135
Fig. 1–13.
Xylocopa coronata Smith 1860: 135; type locality “Kaisaa” (= Kaioa, also spelled Kayoa, Maluku, Indonesia).  
Holotype. Female, labeled “Kaisaa” and “X. coronata Sm., ‘Malay Arch’, F. Sm. coll.” found by the author in collec-
tion of the National Museum of Natural History, Smithsonian Institution, to be returned to the Frederick Smith 
Collection, Oxford University Museum, Fig. 1–4.
Synonyms. Xylocopa combinata Ritsema (1876: 181), type locality “Obi-eilanden” (= Obi, Maluku, Indonesia); 
synonymy by Lieftinck (1956: 63). Xylocopa coronata combinata Ritsema, combination proposed by Lieftinck 
(1956: 63).
Known distribution. Xylocopa coronata coronata Smith: Indonesia: Bacan: Wajaua, Labuha, and unspecified 
localities; Halmahera: Akilamo, Atjengo, Dodinga, Galelo, Goa, Kau, Tolewang, and unspecified localities in 
North, South, and Southeast Halmahera; Kai; Kaioa; Obi: Kasowari and unspecified localities; Ternate: Bukunora 
and unspecified localities; Tidore (Lieftinck 1956: 62). The Kai locality given by Lieftinck (1956: 62) based on old 
specimens in the F. Smith collection requires confirmation, given the significant geographic distance between Kai 
or Kei and the other localities reported here for this species.  Xylocopa coronata combinata Ritsema: Indonesia: 
Obi: Laiwui and unspecified localities (Lieftinck 1956: 63).
Additional material examined. X. coronata coronata: Indonesia: Bacan Island, Kampung Wayamiga, 27–31 July 
1981, A. C. Messer (1 female); “S. Batjan” with no further locality information, VI–VII.1953, A. M. R. Wegner 
(1 female, 1 male). Halmahera Island, Jailolo District, Isthmus of Dodinga, 29 January 1981, A. C. Messer and P. 
M. Taylor (1 female); Jailolo District, Kampung Pasir Putih, 0° 53′ N, 127° 41′ E, 1–14 January 1981, A. C. Messer 
Figures 1–4. Holotype female of Xylocopa coronata and accompanying labels. 1) Dorsal view. 2) Labels. 3) Lateral 
view. 4) Oblique view.
Taxonomy and distribution of Xylocopa coronata Insecta Mundi  0799 · 3
and P. M. Taylor (11 females); same data except 15–31 January 1981 (2 females); same data except 1–14 Febru-
ary 1981 (3 females and 1 male); same data except December 1980 (1 female), same data except 1–14 May 1981 
(2 females); same data except 15–31 July 1981, P. M. Taylor (1 female); same data except no date of collection (1 
female); Kao District, Kao River Basin, Air Kanan, Kampung Tuguls, 1–14 March 1981, A. C. Messer and P. M. 
Taylor (1 female); Kao District, Kampung Kao, 19 March 1981, A. C. Messer and P. M. Taylor (1 female); Tobelo 
District, Kampung Rupo, 11 April 1981, A. C. Messer and P. M. Taylor (1 female). Ternate Island, T. Barbour (1 
female). Tidore Island, Kampung Guaepaji, 5–10 July 1981, A. C. Messer (3 females); same data except 6–7 July 
1981 (8 females). X. coronata combinata: Indonesia: Obi, NW Obi, Laiwui, 0–200 m, IX–X.1953, A. M. R. Wegner 
(2 females); same data except IX.1953 (1 female). All specimens in USNM collection.
Notes on type material of X. coronata. This species was originally described by Frederick Smith from a sin-
gle female specimen collected at “Kaisaa” by Alfred Russel Wallace during his travels in the Malay Archipelago 
(Smith 1860, 1874; see also Lieftinck 1956: 62, who notes that the original description was based on a single speci-
men). According to the detailed chronicle of Wallace’s travels published by Baker (2001), the locality “Kaisaa” 
corresponds to modern-day Kaioa or Kayoa, an island in the Maluku Archipelago of Indonesia. In preparation 
for his revision of the species of Xylocopa from the Maluku Archipelago, Lieftinck (1956: 62) was unable to locate 
any specimens in the Smith collection or other institutional collections which were specifically labeled “Kaisaa,” 
and material from “Kaisaa” is not listed among the numerous specimens of X. coronata that were examined by 
Lieftinck (1956: 62). Instead, Lieftinck (1956: 62) designated a female specimen collected at “Kai” as a lectotype 
for X. coronata. Kai or Kei is an altogether different island group within the Maluku Archipelago which is located 
approximately 872 km from Kaioa (shortest distance according to Google Maps 2020). While examining the 
collection of specimens of X. coronata in the National Museum of Natural History, Smithsonian Institution, I 
discovered a female with a smaller handwritten label stating it was collected at “Kaisaa” and a larger handwritten 
Figures 5–12. Female and male specimens of Xylocopa coronata; all are X. coronata coronata except for the female 
of X. coronata combinata in Figures 8 and 12. 5) Dorsal habitus, female, Halmahera. 6) Dorsal habitus, female, Ba-
can. 7) Dorsal habitus, female, Tidore. 8) Dorsal habitus, female, Obi. 9) Dorsal habitus, male, Bacan. 10) Lateral 
view, male, Bacan. 11) Lateral view, female, Tidore. 12) Lateral view, female, Obi.
4 · September 25, 2020 Mawdsley
label indicating that it was from the F. Smith collection (Fig. 1–4). The specimen corresponds closely with the 
description of the unique holotype provided by Smith (1860: 135) as well as the redescription of the holotype 
given by Smith (1874: 279–280) in his monographic revision of the genus Xylocopa. The accompanying larger 
handwritten label is similar in style and content to labels reported from the F. Smith collection by Lieftinck (1956: 
62), while the handwriting on the smaller label is similar to that of Alfred Russel Wallace in the examples pub-
lished by Baker (2001) and in additional examples of original Wallace labels studied by Mawdsley (2006). This 
specimen had been sent by staff of the Oxford University Museum of Natural History to the late Paul D. Hurd, 
Jr. of the Smithsonian Institution in preparation for his comprehensive monograph reclassification of the genus 
Xylocopa (Hurd and Moure 1963). Based on the evidence available, this specimen appears to be the original holo-
type specimen on which Smith’s description of the species was based.
Notes on geographic variation and subspecies. Lieftinck (1956) placed X. combinata Ritsema 1876 as a sub-
species of X. coronata based on examination of pinned specimens, which included large series of specimens of 
both taxa, the original syntypic series of X. combinata, and multiple specimens of X. coronata which had been 
authoritatively identified by F. Smith. The principal difference between the two forms is in the coloration of the 
pubescence on the dorsal mesosoma in females. In certain populations on the island of Obi, particularly in the 
northwestern portion of the island, females have a broad band of yellow pubescence across the posterior portion 
of the mesosoma (Fig. 8, 12) which is otherwise black in females from elsewhere in the Maluku Archipelago (Fig. 
1, 5–7, 11, 13). This color difference is very consistent in the limited material that I have examined, which sug-
gests a possibility that two separate species may be involved. However, Lieftinck (1956: 64) notes that specimens 
with small amounts of yellow pubescence or individual yellow hairs on the base of the mesosoma occur at sites 
elsewhere in the species range, including the island of Bacan and West Obi. Further studies of morphology, spe-
cies biology, and molecular systematics will be needed to determine the status of the populations on the island of 
Obi. For the time being, I prefer to follow Lieftinck (1956) in recognizing two separate subspecies, which can be 
separated readily by the differences in dorsal coloration of females.
Figure 13. Map showing known distribution of Xylocopa coronata, with schematic illustrations of the dorsal col-
oration patterns in females from each locality.
Taxonomy and distribution of Xylocopa coronata Insecta Mundi  0799 · 5
In my own examination of specimens of X. coronata from the USNM collection, I noticed that the wing 
iridescence in female specimens of this species also exhibits geographic variation. In female specimens identified 
as X. coronata combinata from northwest Obi, the wings are iridescent blue-violet or blue-green, with strong blue 
iridescence predominating (Fig. 8). Female specimens identified as X. coronata coronata from Tidore, Ternate, 
and Kaioa have wing iridescence that is coppery or coppery-violet in color (Fig. 1, 7), while females identified as 
X. coronata coronata from Halmahera and Bacan have wing iridescence that is purplish-blue with strong purple 
iridescence predominating (Fig. 5, 6). Further study of additional specimens is needed to determine the exact 
geographic distribution of these various color variants.  The color of wing iridescence is frequently used as a diag-
nostic character to separate taxa at the species level in other lineages of Xylocopa (see, for example, illustrations 
and discussion in Mawdsley 2017a, 2017b), and thus these different forms clearly merit further investigation in 
order to determine whether multiple valid species-level taxa are present.  However, it should also be noted that at 
least one other widespread species of Xylocopa exhibits infraspecific geographic variation in wing iridescence col-
oration: the Central and South American X. (Neoxylocopa) frontalis (Olivier), in which different populations have 
violet, blue, blue-violet, greenish-blue, golden-green, or dark green wing iridescence, as discussed by Hurd (1978).
Acknowledgments
For permission to examine specimens for this study, I thank Seán Brady and Brian Harris of the National Museum 
of Natural History, Smithsonian Institution (USNM). For critical review and comments on the manuscript prior 
to publication, I thank Sam Droege of the United States Geological Survey’s Bee Inventory and Monitoring Labo-
ratory and Eugene Scarpulla of the Maryland Entomological Society.
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Received June 12, 2020; accepted July 24, 2020. 
Review editor Elijah Talamas.



Notes on the taxonomy and distribution of \u3ci\u3eXylocopa coronata\u3c/i\u3e Smith, 1860, a large carpenter bee from Maluku, Indonesia (Insecta: Hymenoptera: Apidae)

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Notes on the taxonomy and distribution of \u3ci\u3eXylocopa coronata\u3c/i\u3e Smith, 1860, a large carpenter bee from Maluku, Indonesia (Insecta: Hymenoptera: Apidae)

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