A Review of Small Carpenter Bees of the Genus Ceratina , Subgenus ...

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A Review of Small Carpenter Bees of the Genus Ceratina,Subgenus Ceratinidia, of Thailand (Hymenoptera, Apidae)Author(s): Natapot Warrit, Charles D. Michener and Chariya LekprayoonSource: Proceedings of the Entomological Society of Washington,114(3):398-416. 2012.Published By: Entomological Society of WashingtonDOI: http://dx.doi.org/10.4289/0013-8797.1143.398URL: http://www.bioone.org/doi/full/10.4289/0013-8797.1143.398

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PROC. ENTOMOL. SOC. WASH.

114(3), 2012, pp. 398–416

A REVIEW OF SMALL CARPENTER BEES OF THE GENUS CERATINA,SUBGENUS CERATINIDIA, OF THAILAND (HYMENOPTERA, APIDAE)

NATAPOT WARRIT, CHARLES D. MICHENER, AND CHARIYA LEKPRAYOON

(NW, CL) Center of Excellence in Entomology, Department of Biology, Facultyof Sciences, Chulalongkorn University, Bangkok 10330, Thailand; (CDM) Divisionof Entomology, University of Kansas Natural History Museum and EntomologyProgram, Department of Ecology and Evolutionary Biology, University of Kansas,Lawrence, Kansas 66045, USA (e-mail: [email protected]); (NW) Department ofEntomology, National Museum of Natural History, Smithsonian Institution,Washington, D.C. 20013, USA (e-mail: [email protected])

Abstract.—Ten taxa of small carpenter bees belonging to subgenus CeratinidiaCockerell and Porter, 1899, of the genus Ceratina Latreille, 1802, are recorded fromThailand. One new species, C. chiangmaiensis, is described. Ceratina lepida var.sutepensis Cockerell is elevated to full species status as C. sutepensis Cockerell.Ceratina lepida var. sublepida is synonymized under C. sutepensis. Lectotypes aredesignated for Ceratina cognata Smith, C. compacta Smith, and C. sutepensisCockerell. Collecting records and brief taxonomic comments on Ceratinidia speciesin Thailand are presented, together with keys to the species.

Key Words: Xylocopinae, locality records, Thailand, taxonomy, keys

DOI: 10.4289/0013-8797.114.3.398

The genus Ceratina Latreille (Apidae:Ceratinini), consists of 23 subgenera andis found on all continents (Terzo 2000).None of the subgenera occur naturallyin both the Eastern and Western hemi-spheres. Eight subgenera, includingCeratinidia, contain Oriental species. ThesubgenusCeratinidiaCockerell and Porter,1899, consists of small to medium-sizednonmetallic black bees with yellow mac-ulations; these are usually solitary and stemnesting. Its range covers major parts ofAsia—Afghanistan to Japan (west to east)and the Primorskyi region of Russia toIndonesia and western New Guinea (north

to south). Habitats include cultivated areaswhere the original vegetation has entirelydisappeared, often in the neighborhood ofhuman dwellings in towns and villages, inaddition to less disturbed areas.

The most recent revisionary work onthe systematics of the Oriental species ofCeratina was by J. van der Vecht (1952).In this work, 22 species of Ceratinidiawere recognized, seven of which wererecorded from Thailand—C. accusatorCockerell, C. bryanti Cockerell, C.cognata Smith, C. collusor Cockerell, C.compacta Smith, C. lieftincki van derVecht, and C. nigrolateralis Cockerell.Van der Vecht (1952) recognized twospecies groups in Ceratinidia, basedprimarily on characters of the male sixth*Accepted by Michael W. Gates

sternum (S6) and seventh tergum (T7)—the bryanti and the compacta speciesgroups. A third species group (the fla-vipes species group) was proposed laterby Shiokawa and Hirashima (1982); it isbest known from East Asia, e.g., Japan,Korea, and eastern China. The subapicaldepression of the male S6 in the bryantispecies group is equipped with twosubmedian teeth, often with a smalltooth between them (Fig. 4), whereasa distinct median V- shaped ridge (Fig. 3)is present on the subapical depression ofthe species of the compacta species group.In the flavipes species group, the subapicaldepression is equipped with two longsubmedian teeth that are slightly curvedoutward apically and several times longerthan wide (Shiokawa 1963, Shiokawa andHirashima 1982). The flavipes speciesgroup is not recorded from Thailand.

This paper is modified from an as yetunpublished revision of the entire sub-genus (Warrit 2007a). Ceratinidia spec-imens from Thailand deposited in elevenmajor museums have been studied, as wellas material from a collecting excursion byNW to Thailand in 2003. Ten species arerecognized (including one form, C. in-certula, that is of uncertain status). Allseven species that van der Vecht previouslyrecorded from Thailand are included. Onenew species is described, and one name israised to specific status. Taxonomic com-ments on the known and newly describedspecies and identification keys to bothsexes are provided. A recent study (Warrit2007a) shows thatCeratinidiamay later beexpanded to include the subgenera Lio-ceratina van der Vecht and Xanthoceratinavan der Vecht, but this paper does not in-clude species of those subgenera.

MATERIALS AND METHODS

A total of 120 specimens from Thailandused in this study were borrowed from 11

institutions. These institutions are listedbelow. An additional 401 specimens werecollected (into ethanol) by NWin Thailandfrom June to August, 2003, for a total of521 specimens from Thailand used in thisstudy.

Specimens were examined and mea-sured with an Olympus SZ60 dissectingmicroscope equipped with an ocularmicrometer or calipers. Photomicro-graphs were taken using EntovisionPhotographic System� and MicroOpticsDigital Image Systems�. Illustrationswere prepared using Adobe Photoshop7.0� and Adobe Illustrator 10.0�. Dis-sected male terminalia (S6, T7, andgenitalic structures) were cleared usingca. 3M potassium hydroxide (KOH) inwater at room temperature and stored inglycerin in microvials on pins with dryspecimens. Label data for all specimenswere completely recorded. Such datafor primary type material and paratypesare presented exactly as given on thelabels.

Many general terms for the externalmorphological features used here arebased upon studies of the honeybee(Apis mellifera) by Snodgrass (1956)and other bees by Michener (1944, 2000).Taxon-specific terminology for Ceratinafollows van der Vecht (1952), Daly (1973),and Terzo (2000).

We have tried to examine the name-bearing types for all species and haveindicated for each species whether wehave seen its type, and in what collectionthe type is found. When unable to exam-ine a name-bearing type, we indicate thebasis for our recognition of the name—examination of a paratype, specimensidentified by the author of the name, ormerely the printed information. If anolder author stated with his descriptionthat he had only a single specimen, weregard it as the holotype, even if it is onlylabeled “Type”.

VOLUME 114, NUMBER 3 399

Some authors such as F. Smith (1854,1879) often did not say upon how manyspecimens they based a description.Commonly one specimen is found, la-beled as the TYPE. In such cases wedesignate that specimen as the lectotype(since designation by unpublished la-beling is not decisive). We do this inaccordance with ICZN, fourth edition,Article 74.5 so that if additional speci-mens are found later in the same or an-other collection, none of them would beeligible for lectotype designation. Thus theusage of the name would not be subject topossible instability.

Theworks of T. D. A. Cockerell presentspecial problems for understanding oftypes. Except perhaps in his very early(before 1900) works, Cockerell normallylabeled, for each new species, one speci-men as TYPE and the others, if any in thetype series, as COTYPES (CDM, pers.comm.). In current terminology, the TYPEfor Cockerell would now be consideredthe holotype while his COTYPES wouldnow be considered paratypes. We haverespected these usages, considering hisTYPE as holotype, as indicated by ICZN,fourth edition, Article 74.5.

Acronyms of Institutions

BKOK Chulalongkorn University Museumof Natural History, Bangkok, Thailand

IZCAS Academia Sinica (Chinese Acad-emy of Sciences), Beijing, China

CMAI Chiang Mai University Entomolog-ical Collection, Chiang Mai, Thailand

BPBMBishopMuseum, Honolulu, Hawaii,USA

SEMCUniversity of Kansas Natural HistoryMuseum and Biodiversity Research Center,Lawrence, Kansas USA

RMNH National Natuurhistorisch Museum,Leiden, The Netherlands

BMNH The Natural History Museum,London, United Kingdom

LACM Los Angeles County Museum of Nat-ural History, Los Angeles, California, USA

AMNH American Museum of NaturalHistory, New York City, New York, USA

CAS California Academy of Sciences, SanFrancisco, California, USA

USNM National Museum of Natural His-tory, Smithsonian Institution, Washington,D. C., USA

These acronyms are used to indicatethe locations of specimens in the accountof each species, except that for the largenumber of specimens collected by NW,no acronym is given. These specimensare in alcohol, not in the pinned collec-tion, at SEMC.

RESULTS AND DISCUSSION

Subgenus Ceratinidia Cockerelland Porter

Ceratina (Ceratinidia) Cockerell andPorter, 1899: 406. Type species:Ceratinahieroglyphica Smith, 1854, by originaldesignation.

Diagnosis.—Ceratinidia has, on alltagmata, extensive yellow maculationon a dark (usually black) nonmetallicbackground. The frons and vertex arerather strongly punctate; punctures existon the paraocular area above the levelof the antennal socket and in the spacebetween the compound eye and theocelli. Males have a tuft of long hairs,usually divided into two parts, on theapex of the gonostylus. Two other sub-genera, Lioceratina van der Vecht, 1952,and Xanthoceratina van der Vecht, 1952,have more extensive yellow maculationon the clypeus, paraocular area, scutum,mesepisternum, scutellum, metanotum,and terga; they thus appear largely yellow.They also have more delicate, less densepunctuation and lack the hair tufts on themale gonostyli. These subgeneramay laterbe included in Ceratinidia (Warrit 2007a).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON400

Key to Species of Ceratinidia ofThailand (Females)

1 Area between lateral margin of scutum and

parapsidal furrow (Fig. 1) shiny, im-punctate, or with, at most, one incomplete

row of punctures along outer side of para-psidal furrow . . . . . . . . . . . . . . . . . . . . 2

— Area between lateral margin of scutum and

parapsidal furrow punctate with one orusually two or more complete rows of

punctures along outer side of parapsidalfurrow . . . . . . . . . . . . . . . . . . . . . . . . . 8

2(1) Procoxa angularly produced laterally asprominent process (Fig. 2) . . . . . . . . . . 3

— Procoxa rounded on outer side, withoutprominent process . . . . . . . . . . . . . . . . 4

3(2) Frontal area above frontal spot sparselypunctate, with no more than 15–20 punc-

tures; area between compound eye anddorsolateral margin of antennal fossa im-

punctate (at most with 1 or 2 punctures);base of antennal scape often yellow or fer-

ruginous, with abrupt color change, some-times apex yellow; marking on paraocular

area gradually tapering upward (as in Fig. 6);mesotibia dark brown to black with yellow

line on outer side usually extending for entiretibial length . . . . . . . . . . . . . . . C. collusor

— Frontal area above frontal spot densely punc-tate, punctures occupying almost entire area;

area between compound eye and dorsolateralmargin of antennal fossa varied (densely

punctate, sparsely punctate, with few punc-tures, or impunctate); base of antennal

scape ferruginous to black, apex black;marking on paraocular area varied (gradu-

ally tapering upward or widened abruptly atmedian level from top to bottom); marking on

outer side of mesotibia varied (absent, smallspot at base, or yellow line extending for en-

tire tibial length) . . . . . . .C. nigrolateralis4(2) Punctures on mesepisternum widely spaced,

seperated by 1–1.5 times puncture diam-

eters, posterolateral area of mesepisternumnear mesocoxal base with shiny area

with few punctures (Fig. 7); hypoepimeralarea with large impunctate space above

scobe . . . . . . . . . . C. chiangmaiensis n. sp.— Punctures on mesepisternum dense, inter-

spaces less than puncture diameter, postero-lateral area of mesepisternum near mesocoxal

base with punctuation only slightly less densethan lateral and ventral areas; hypoepimeral

area with or without large impunctate areaabove scrobe . . . . . . . . . . . . . . . . . . . . 5

5(4) Paraocular area with dense small to coarse

punctures clumping together on medianpart, few punctures on lower part; clypeus

with conspicuous median longitudinal im-pressed area one third as wide as upper

lobe of clypeus, dorsal and dorsolateralarea of clypeus with dense small to me-

dium-sized punctures, median area usually

smooth and impunctate, but sometimes

slightly rugose with few punctures . . . . 6

— Paraocular area without clump of dense

punctures on median part, pattern of

punctures varied, with few ill-defined

punctures or with scattered punctures

but interspaces between punctures broader

than puncture diameters; clypeus smooth,

almost impunctate or with small fine

punctures along dorsal and dorsolateral

margin . . . . . . . . . . . . . . . . . . . . . . . . 76(5) Scutum with four longitudinal yellow lines

(Fig. 1) or the submedian lines sometimesweak or even absent; yellow transverse band

on pronotum usually unbroken; markingson pronotal lobe and scutellum usually

present . . . . . . . . . . . . . . . . C. bryanti

— Scutum without longitudinal yellow lines;yellow transverse band on pronotum

often interrupted medially; markings onpronotal lobe and scutellum usually

absent. . . . . . . . . . . . . . . . . . . . .C. sutepensis7(5))Upper lobe of inverted-T mark on clypeus

well-developed, at least reaching 2/3 ofclypeal length to almost attaining fronto-

clypeal sulcus (apex more or less truncate),base of antennal scape usually yellow;

metasomal bands well developed and un-broken (or T3 band interrupted), except no

band on T6; T1 with wide band enclosingtwo small black spots . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . C. collusor (part)— Upper lobe of inverted-T mark on clypeus

small, reaching at most half of clypeallength, or absent (apex sometimes

pointed); base of antennal scape ferrugi-nous or black; T1 band usually reduced to

small lateral spot or absent; T2 and T3 bandsbroadly interrupted medially, usually abruptly

broadened laterally . . . . . . . . . . . C. accusator8(1) Scutum with four longitudinal yellow lines;

labrum sometimes with yellow spot . . . 9— Scutum without longitudinal yellow lines;

labrum always black, without yellowspot . . . . . . . . . . . . . . . . . . . C. cognata

9(8) Base of antennal scape yellow; posterolat-eral area of mesepisternum near mesocoxal


base densely punctate (interspaces smaller

than puncture diameters) . . . . . . . . . . . . . . .

. . . . . . C. compacta (see also C. incertula)

— Base of antennal scape black; posterolateral

area of mesepisternum near mesocoxal base

sparsely punctate (interspaces broader than

puncture diameters) . . . . . . . . C. lieftincki

Key to Species of Ceratinidia inThailand (Males)

(Males are unknown for C. chiang-maiensis and C. incertula)

1 Subapical depression of S6 with distinctmedian V-shaped ridge (Fig. 3), often with

small tooth on each side; small projection

between teeth absent (compacta species

group) . . . . . . . . . . . . . . . . . . . . . . . . . 2

— Subapical depression of S6 without median

V-shaped ridge, with two submedian teeth

and often with small median tooth between

the submedian teeth (Fig. 4) (bryanti species

group) . . . . . . . . . . . . . . . . . . . . . . . . . 6

2(1) Area between the lateral margin of thescutum and parapsidal furrow (Fig. 1)

shiny, impunctate, or with, at most, one

incomplete row of punctures along the

outer side of parapsidal furrow . . . . . . . 3

— Area between the lateral margin of the scutum

and parapsidal furrow with one or usually two

or more complete rows of punctures along the

outer side of parapsidal furrow . . . . . . . . 43(2) Frontal area above frontal spot sparsely

punctate, with no more than 15–20 punc-

tures; area between compound eye and

dorsolateral margin of antennal fossa shiny

and impunctate (at most with 1 or 2 punc-

tures); base of antennal scape often yellow

or ferruginous, with abrupt color change,

sometimes apex yellow; yellow marking on

mandible sometimes present . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . C. collusor

— Frontal area above frontal spot densely

punctate, punctures occupying almost entire

area; area between compound eye and dor-

solateral margin of antennal fossa varied

(densely punctate, sparsely punctate, with

few punctures, or impunctate); base of an-

tennal scape ferruginous to black, apex al-

ways black; mandible dark brown or black

. . . . . . . . . . . . . . . . . . . C. nigrolateralis4(2) Scutum with four longitudinal yellow lines

(as in Fig. 1); yellow spot behind pronotal

lobe distinct . . . . . . . . . . . . . . . . . . . . . 5

— Scutum without longitudinal lines (some-

times with two vague outer lines); yellow

spot behind pronotal lobe often absent or

small (sometimes distinct) . . . C. cognata

5(4) Base of antennal scape yellow . . . . . . .. . . . . . . . . . . . . . . . . . . C. compacta

— Base of antennal scape black or with

vague yellow spot . . . . . C. lieftincki6(1) Paraocular area smooth with sparse fine

punctures (interspaces broader than 2 times

puncture diameters); clypeus usually

smooth and shiny with scattered fine punc-

tures, mostly along upper margin (some-

times upper portion slightly rugose); scutum

with four longitudinal yellow lines; marking

behind pronotal lobe usually present; area

between parapsidal furrow and lateral mar-

gin of scutum shiny and impunctate; body

length 6.5–7.3 mm . . . . . . . . C. accusator

— Paraocular area more or less rugose with fine

and coarse punctures, usually concentrated

in median area; unmarked area on upper

portion of clypeus strongly rugose or at least

with dense coarse punctures; scutum without

longitudinal yellow lines (sometimes with

two vague outer lines); marking behind

pronotal lobe absent; area between para-

psidal furrow and lateral margin of scu-

tum usually with punctures; body length

7.0–9.6 mm . . . . . . . . . . . . . . . . . . . . 7

7(6) Pronotal lobe, scutellum, hind tibia, andterga with yellow markings. . .C. bryanti

— Pronotal lobe, scutellum, hind tibia, and

terga without yellow markings . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . C. sutepensis

Ceratina accusator Cockerell

Ceratina accusator Cockerell, 1919a: 249[female and male]; Cockerell, 1920a: 624.van der Vecht, 1952: 56–57 [note on ho-lotype and additional material Ceratinamentioned in key]; Cockerell, 1929: 150[record in Nan Province, Thailand].

Type Material.—The female holotypeof C. accusator in AMNH is labeled“Ceratina accusator Ckll.”, “Island ofPenang, Baker” [Penang Island, Malaysia],and “acc. 34970”.

Thai Material Examined.—Chumpon:Lang Suan District, 15 July 2007, 10° 019N99° 039E, N. Warrit (4♀, 2♂). Prachup


Kirikhan: Meuang District, 26 June 2003,11° 409N 99° 419E, N. Warrit (3♀). SamutSakorn: Mae Klong District, 3 July 2003,N. Warrit (2♀, 1♂).

Distribution.—Ceratina accusator isknown from southern Thailand, Malaysia,and Indonesia. The distribution may reachnorthern Thailand, since a specimen fromNan [Thailand] was recorded by Cockerell(1929). We have not seen this speci-men but given its locality, it could beC. chiangmaiensis.

Diagnosis and comparative comments.—Ceratina accusator is a small speciescompared to most other Ceratinidia(6.20–6.80 mm). In the female, the upperlobe of the marking on the clypeus is poorlydeveloped, reaching at most half of the cly-

peal length. The marking on the paraoculararea is short, usually reaching at most the

level of the antennal socket. In both sexes,

many specimens have the yellow bands

widely interrupted medially on T2 and T3.Ceratina accusator is similar to C.

chiangmaiensis in body size and yellow

markings. However, C. accusator can be

distinguished from the later by the dense,

well-defined punctures on the clypeus,

paraocular area, frons, vertex, anterior

portion of scutum, and mesepisternum.

Ceratina bryanti Cockerell(Figs. 4, 10, 11)

Ceratina bryanti Cockerell, 1919b: 175[male]; van der Vecht, 1952: 51–54

Figs. 1–4. Diagram of dorsum of thorax of a Ceratinidia. Gray areas represent yellow maculations.Fig. 2. Diagram of foreleg of female Ceratinidia showing coxal process of some species. Figs. 3–4. Sixth

sternum of male of Ceratina compacta Smith and C. bryanti Cockerell, respectively.


[redescription]; Baker, 2002: 361, fig. 5[indicated synonyms].

? Ceratina denticulata Wu, 1963: 86, 91, fig.82 [female and male; so far as can be judgedfrom the disintegrated allotype and fromWu’s original description, C. denticulatamay be a synonym of C. bryanti or C. hi-eroglyphica Smith].

Type Material.—The male holotypeof C. bryanti in USNM is labeled “Pe-laboean Ratoe Java” “Bryant & PalmerColl.”, and “Type No. 20706 U.S.N.M.”.

Themale holotype ofC. denticulata is inIZCAS (not seen). The female allotypein the same institution is labeled “[localityin Chinese], 200 m., 1956. VI. 11”,“Allotype”, “IOZ (E) 206055”, and“Ceratina denticulata Wu f#”. The allo-type was in poor condition when it arrivedat the University of Kansas in 2007 forexamination. The head and metasoma fellapart from the mesosoma, the head wasmissing from inside the container con-taining the type, and most parts of the legssegments were broken from themesosoma.

Thai Material Examined.—NakornRatchasrima: Suranaree TechnologicalUniversity, 20 June 2003, 14° 539N102° 009E, N. Warrit (1♀). PrachupKirikhan: Bang Sapan District, 26 June2003, 11° 209N 99° 299E, N. Warrit(15♀); Pran Buri District, 27 June 2003,12° 229N 99° 539E, N. Warrit (1♀).Ranong: Kuraburi District, 16 July 2003,09° 199N 98° 259E, N. Warrit (14♂).

Distribution.—Ceratina bryanti isknown from southern Thailand, Malaysiaand some of the Indonesian Islands (e.g.,Sumatra, Java, and Bali). Ceratina den-ticulata is known from Southern China.

Remarks.—Based onWu’s description(1963) and illustrations of C. denticulata,it is somewhat convincing to think thatC. denticulata is a synonym of C. bryantior C. hieroglyphica sensu van der Vecht(1952). The marking on the clypeus and

paraocular area below the antennal fossa,the submedian teeth on S6 (Wu’s illus-tration did not indicate the presence ofa small median denticle between thesubmedian teeth), and the form of T7 ofthe male of C. denticulata are similar tothose of C. bryanti and C. hieroglyphica.However, markings on the pronotum,scutum, and legs of the female allotype ofC. denticulata are absent or more reducedthan in material of the other two species,suggesting that C. denticulata may be adifferent species.

Ceratina chiangmaiensis Warrit,Michener, and Lekprayoon, new species

(Figs. 5–7)

Description.—Female: Structure: Length5.15–6.00 mm. Prosoma: Clypeus smoothwith scattered shallow punctures, mostlyon upper and lateral margins; medianlongitudinal depressed area with vaguelongitudinal carina. Labrum with mixtureof fine and coarse punctures, and withlong bristles. Paraocular area below an-tennal fossa shiny, almost impunctate,with shallow punctures along compoundeye. Inner part of antennal fossa withrather dense punctures along frontal ca-rina, outer part impunctate. Supraclypealarea below frontal carina impunctate.Frons with few punctures; vertex behindocelli with denser and coarser punctures.Space between ocelli and upper part ofcompound eye with few punctures. Genashiny, almost impunctate. Preoccipitalcarina not prominent; area behind ocellislightly depressed. Hypostomal area slightlydepressed (sometimes rugose). Mesosoma:Anterior third of scutumwith fine punctures(interspaces broader than puncture di-ameters); posterior two thirds of scutumshiny and almost impunctate except pos-terior margin; few punctures along an-teromedian line and parapsidal furrow.Lateral margin of scutum lined with one or


two rows of punctures. Procoxa withrounded angle on outer side at base. Me-sepisternum with punctures not as dense asin C. accusator; punctures noticeably lessdense ventrally and posterolaterally near

mesocoxal base. Propodeal triangle finelycoriaceo-reticulate; lateral area of propo-deum densely and finely punctate, withsmall patch of dense hair anteriorly. Mac-ulation (yellow except as noted): Prosoma:

Figs. 5–7. Ceratina (Ceratinidia) chiangmaiensis n. sp., holotype female, lateral view, face, and

mesepisternum. Fa, Frontal area; Pa, Paraocular area; Im, Inverted T-shaped mark of clypeus; Ha, Hy-poepimeral area; Pl, posterior lateral area of mesepisterum.


Upper lobe of inverted-T marking ofclypeus reaching half clypeal length(sometimes with small midapical incisionon upper margin of marking). Labrumand mandible black. Supraclypeal andfrontal spots well developed. Marking onparaocular area linear, tapering upward(Fig. 6). Genal marking slightly narroweddownward; extending half of eye length.Mesosoma: Transverse marking on pro-notum interrupted medially and rarelyconnected to yellow spot on pronotallobe. Spot behind pronotal lobe variable.Scutum with four longitudinal lines. Ax-illa black. Tegula reddish-brown trans-lucent. Scutellum with small triangularmarking, anterior margin with slight in-cision. Marking on metanotum absent.Protrochanter and mesotrochanter withsmall yellow marking apically. Profemurwith yellow spot on outer side at apex.Protibia and mesotibia black with yellowmarking on outer side extending almostentire tibial length. Metatibia with yellowspot at base. Tarsi dark brown except met-atarsi yellow brown.Metasoma:Metasomaltergal bands narrow but well developed andunbroken, T2 and T3 bands abruptly wid-ened laterally; no band on T6; T1 bandwide laterally, not enclosing black spots.

Male: Unknown.Type Material.—The female holotype

and two paratypes are in SEMC. Theyare in excellent condition and are labeled“THAILAND: Chiang Mai Prov., AhnKang, W. of Fang, 6 February 1993,S. Boongird, C. Michener, S. Malaipan”.Three additional paratypes with the samedata are deposited, one each, in BMNH;USNM; and BKOK.

The type material was taken from nestburrows in broken or cut dead stems ofLantana sp. (Family Verbenaceae). Noimmature stages or provisioned cells werefound; the active season must have beenat another time of year (CDM, personalobservation).

Etymology.—This species is namedafter Chiang Mai Province, Thailand,where all of the types were collected.

Distribution.—Ceratina chiangmaien-sis is known only from the type locality inthe mountains of Chiang Mai Province,Thailand.

Diagnosis and Comparative Com-ments.—Ceratina chiangmaiensis is sim-ilar to C. accusator, but is even smallerthan the later species (5.15–6.00 mm inbody length). A character that separatesC. chiangmaiensis from C. accusator isthe reduction of punctures, mostly on thefacial area and the scutum. The para-ocular area is smooth, shiny, and some-times impunctate; the frons is sparselypunctate with shallow punctures; theclypeus is not as punctate and rugose as inC. accusator; the anterior part of thescutum is sparsely punctate with shallowpunctures; and the mesepisternum issparsely punctate (interspaces betweenpunctures are larger than the puncturediameter).The yellow maculation behindthe pronotal lobe is sometimes absent.

Ceratina cognata Smith

Ceratina cognata Smith, 1879: 94 [male]; vander Vecht, 1952: 69–73 [redescription of fe-male and male]; Wu, 2000: 168–169 [recordfrom Southern China].

Ceratina conscripta Cockerell, 1919a: 247[male]; Cockerell, 1920a: 624 [mentioned inkey]. Synonymy by van der Vecht (1952).

Ceratina selangorensis Cockerell, 1919a: 248[male]. Synonymy by van der Vecht(1952).

Ceratina laosorum Cockerell, 1929: 151 [fe-male]. Synonymy by van der Vecht (1952).

Type Material.—The male type of C.cognata in BMNH, is labeled “Celebes”[Sulawesi Island, Indonesia], “57 101”,and “Type # 17 b 213”. This specimen isin good condition. Since Smith (1878)and subsequent authors did not indicate


how many specimens Smith had, nor didthey designate a lectotype, the male syn-type of C. cognata as labeled in BMNH ishere designated as lectotype with a redlabel “Ceratina cognata Smith, Lecto-type, N. Warrit, 2007”.

The male holotype of C. conscriptais in BMNH (not seen). However, vander Vecht (1952) made a note that al-though this type is from Penang, Malaysia,the color pattern agreed well with thetype of C. cognata from Celebes (Sula-wesi). Cockerell (1919a) reported thelabel on the type as “Island of Penang(Baker, 9286)” at the end of the speciesdescription.

The male type of C. selangorensis inBMNH, is labeled “Selangor Baker”[Selangor, Malaysia], “9287”, and“Type # 17 b 237”. It is in good con-dition; however, the yellow maculationsare yellowish red, presumably from theprolonged use of cyanide as the killingagent. The type of C. selangorensis inBMNH is here designated as lectotypewith a red label “Ceratina selangorensisCockerell, Lectotype, N. Warrit, 2007”.NW has examined this lectotype andagrees with van der Vecht’s (1952) com-ment that it is a dark form of C. cognata.The punctation and yellow maculation ofC. selangorensis are as in C. cognata,except that in C. selangorensis thetransverse yellow bands on T2–T4 arevague spots, and the remaining segmentsare black.

The female holotype of C. laosorumin BMNH, is labeled “Brit. Mus. 1933–567” and “Type # 17 b 223”. The type isin good condition and falls within therange of variation of C. cognata; it is asmaller specimen with extensive macu-lation on the tibiae and tarsi. As ex-plained in Materials and Methods, it isthe holotype.

Thai Material Examined.—7 April1973, on flowers of Ziziphus, P.S. Mes-

senger (1♀; CAS), 19 May 1973, onflowers, P.S. Messenger (8♀, 1♂; CAS),6 June 1973, on flowers, P.S. Messenger(2♀; CAS). Chantaburi: ; 2 km. S. —Bangkok of: 1 Chantaburi, 19–21 June1969, 12° 369N 102° 079E, J.J.S. Burton(1♀; BPBM). Chiang Mai: Amphur MaeWang, Thanbon Donpao, Pa Huay Kho,20–30 April 1997, 18° 41.549N 98°48.4179E, 250 m., Malaise trap in fieldat edge of deciduous forest, SaowapaSonthichai (1♀; LACM). Chiang Rai:Meuang District, 12 June 2003, 20° 059N99° 519E, N. Warrit (1♀); Rajabhat In-stitute, 13 June 2003, N. Warrit (3♀).Chumpon: Meuang District, 25 June2003, 10° 339N 99° 079E, N. Warrit (1♀,2♂); Sapree District, 25 June 2003, 10°349N 99° 169E, N. Warrit (1♀); SaweeDistrict, 15 July 2003, 10° 209N 99°059E, N. Warrit (3♂); Tha Sae District,25 June 2003, 10° 529N 99° 169E, N.Warrit (2♀, 3♂). Kanchanaburi: 110 airkm. W. of Bangkok, 25–27 April 1988,W.J. Pulawski (2♀; CAS). NakhonNayok: Nakhon Nayok, 21 January1993, S. Boongird and C.D. Michener(2♀, 1♂; SEMC). Nakhon Ratchasima:Suranaree Technological University, 20June 2003, 14° 539N 102° 009E, N.Warrit (11♀); Tha Chang, 10 February1993, ex. Callistemon, S. Boongird andC.D. Michener (1♀; SEMC). Nakhon SriThammarat: Chalerm Prakiet District, 18July 2003, 08° 109N 100° 029E, N. Warrit(3♀, 2♂). Phuket: Phuket Island, 29January 1993, S. Boongird and C.D.Michener (2♀; SEMC). Prachup Kirikhan:Kui Buri District, 27 June 2003, 12°059N 99° 509E, N. Warrit (4♀). Ranong:Kuraburi District, 16 July 2003, 09° 199N98° 259E, N. Warrit (1♀, 14♂). Saraburi:August, Sakda (in Thai) (1♀; BKOK).Surat Thani: Bang-O 25 km. SW. of SuratThani, 30 January 1993, S. Boongird andC.D. Michener (1♀; SEMC); Ko SamuiDistrict, 17 July 2003, 09° 279N 99° 569E,


N. Warrit (3♂). Trang: Khao ChongNature Education Center, 21–24 July1996, 07° 359N 99° 469E, Snelling andSonthichai (3♀; LACM); Sikoa District,19 July 2003, 07° 319N 99° 239E, N.Warrit (1♀).

Distribution.—This widely distributedspecies occurs from southern China, theIndochinese region (including Thailand),to the Malay Peninsula and Indonesia(including Sulawesi), but it is not found inthe Philippines.

Diagnosis and Comparative Com-ments.—Ceratina cognata closely re-sembles C. compacta. Punctation in C.cognata is very similar to that of C.compacta in density, puncture size, anddistribution, with the exception of thedensity of punctures on the postero-lateral mesepisternal area near themesocoxal base. In C. cognata, thesepunctures are widely separated and welldefined; but in C. compacta the puncturesare dense and the surface dull (this char-acter is more evident in most femalesthan in males). Color patterns of the twospecies are also similar; however, C.cognata can be easily distinguished fromC. compacta by the absence of yellowmaculation on the labrum (only in fe-male) and mesoscutum. S6 of male C.cognata has a V-shaped ridge and a smalllateral tooth on each side of the subapicaldepression.

Ceratina collusor Cockerell

Ceratina collusor Cockerell, 1919a: 248[male]; Cockerell, 1920b: 227 [C. collusorvar. a., female and male]; van der Vecht,1952: 75–77 [redescription of female andmale]

Type Material.—The male holotypein BMNH is labeled “T.D.A. CockerellB.M. 1936-415”, “1879b”, “SingaporeColl. Baker”, and “Type # 17 b 238”.The type is in good condition.

Thai Material Examined.—ChiangMai: Chiang Dao District, 9 June 2003,19° 269N 98° 579E, N. Warrit (6♂); MaeRim, 27 January 1994, R.A. Beaver (1♀;CAS); Queen Sirikit Botanical Garden,11 June 2003, 18° 539N 98° 519E, N.Warrit (4♀, 1♂). Chumpon: Chai YaDistrict, 14 July 2003, 09° 199N 99°089E, N. Warrit (2♂). Nakhonsri Tham-marat: Thasala, 24 June 2001, 8° 469N99° 559E, N. Warrit (1♂). Phetburi:Khao Yoi District, 24 June 2003, 13°159N 99° 479E, N. Warrit (2♀). Phuket:29 January 1993, S. Boongird and C.D.Michener (1♀; SEMC). Ranong: KuraburiDistrict, 16 July 2003, 09° 199N 98°259E, N. Warrit (9♀, 9♂). Trang: SikoaDistrict, 19 July 2003, 07° 319N 99°239E, N. Warrit (1♂)

Distribution.—Ceratina collusor isrecorded from Burma, Thailand, Laos,Malaysia, Singapore, and Indonesia.

Diagnosis and Comparative Com-ments.—Both sexes of C. collusor can bedistinguished from otherCeratinidia in thecompacta species group by the absence ofpunctures along the parapsidal furrow andin the area between the furrow and thelateral margin of the scutum, and usuallyby the presence of a protuberance on theouter side of the procoxa. Ceratina col-lusor is similar to C. nigrolateralis inmany aspects, but can be differentiated bythe smaller size, the presence of yellowmarking on the base of the scape (some-times on the apex also), and the rectan-gular marking that occupies most of thescutellum.

Van der Vecht (1952) synonymized C.incertula under C. collusor without ex-amining the type and concluded that it isa dark form of C. collusor. We examinedthe type of C. incertula and found that itis a different species from C. collusor, asshown by the presence in C. incertula ofpunctures along the parapsidal furrowsand in the area between the furrows and


the lateral margin of the scutum, and bythe absence of a protuberance on theouter side of the procoxa.

Ceratina compacta Smith

Ceratina hieroglyphica Smith, 1854: 226[female and male description, part (Phil-ippine specimen)].

Ceratina compacta Smith, 1879: 91 [male;erroneously described as female]; van derVecht, 1952: 66–69 [recognized C. philip-pinensis = C. compacta; redescription offemale and male]; ? Wu, 2000: 172 [recordfrom China; however, only one specimen];Warrit, 2007b: 72–77 [probably adventivein Thailand].

Ceratina philippinensis Ashmead, 1904: 2 [Fe-male]; Cockerell, 1915: 109 [record in Ne-gros, Philippines]; Cockerell, 1916: 304[suspected C. philippinensis = C. compacta];Cockerell, 1918: 142 [records in Mindanaoand Palawan, Philippines]; Cockerell,1920c: 146 [records in Culasi and BatbatanIsland, Philippines]; Cockerell, 1920d: 207;Cockerell, 1925a: 54 [record in Bangui,Ilocos Norte, Philippines]; Cockerell,1925b: 172 [record on Samar Island,Philippines]; Hedicke, 1926: 419. Synonymyby van der Vecht (1952).

Type Material.—The male type of C.compacta in BMNH, is labeled “Phil.Isla.” [Philippines] and “Type # 17 b 214”.This specimen was first described underthe name C. hieroglyphica Smith alongwith material from northern India andHong Kong. Smith (1879) recognizedlater that the specimen(s) from the Phil-ippines differ from those from the twoother locations, and named the form fromthe Philippines as C. compacta. As ex-plained in Materials and Methods, sincethere may have been more syntypes, themale type of C. compacta in BMNH ishere designated as the lectotype and car-ries a red label “Ceratina compactaSmith, Lectotype, N. Warrit, 2008”.

Ashmead’s description of C. philip-pinensis was based on four specimens,one of which was marked as the type.It must be regarded as the holotype andis a female in USNM labeled “ManilaPI” [Manila, Philippines], “Type No7692 U.S.N.M.”, and “W. A. StantonCollector”. According to van der Vecht(1952), specimens of C. compacta havelong been identified as C. philippinensis.This is partly due to the fact that the typeof C. compacta was first erroneouslydescribed as female, instead of male,by Smith (1879). Cockerell (1916) sus-pected that the female of C. philippinensiswas possibly a female C. compacta;later in the same paper he made a noteabout it. Van der Vecht (1952) exam-ined the types of both C. philippinensisand C. compacta and verified thatC. philippinensis is in fact the femaleof C. compacta. NW has seen bothtype specimens and agrees with vander Vecht’s finding.

As noted in the account of C. in-certula Cockerell, that name is a possi-ble synonym of C. compacta, and wouldbe a name for the Thailand populationsof this species if needed.

Thai Material Examined.—Bangkok:Kampengsan district, 10 December 2002,P. Angmani (1♀; SEMC). Chiang Mai:Mae Rim 18° 089N 98° 979E, 13 July1996, R.R. Snelling (1♀; LACM);Meuang District, Mae Hei ResearchStation 18° 459N, 98° 569E, 9 June 2003,500 m., N. Warrit (5♀). Kanchanaburi:100 air km W of Bangkok, 25–27 April1988, W.J. Pulawski (1♂; CAS). Loei:Wang Saphung District, 20 April 1989,W.J. Pulawski (1♀; CAS), 23 April 1989,W.J. Pulawski (1♀; CAS). NakornNayok: 21 January 1993, S. Boongird andC. Michener (1♀; SEMC). Phitsanulok:Bangraka District, Naresuan University16° 449N 100° 119E, 17 June 2003, N.Warrit (1♀).


Distribution.—In earlier literature(van der Vecht,1952), Ceratina compactais recorded only from the Philippine Is-lands. Examination of specimens fromThailand and one from northern Sulawesidocument its presence outside the Philip-pines. The specimens from Thailand mayrepresent a population introduced into thatarea (evidence in Warrit, 2007b), but seealso the account of C. incertula below.

Diagnosis and Comparative Com-ments.—The male of C. compacta hasa V-shaped ridge on the subapical de-pression of S6 instead of submedian teethas in the bryanti species group. Bothsexes of C. compacta can be distin-guished from other Ceratinidia in thecompacta species group by the followingcombination of characters: (1) lateral areaof scutum with one or usually two ormore complete rows of punctures alongthe parapsidal furrow; area betweenlateral margin of scutum and parapsidalfurrow densely or partially punctate; (2)scutum with four longitudinal yellowlines; and (3) base of antennal scapeyellow.

Ceratina incertula Cockerell

Ceratina incertula Cockerell, 1937: 12[female]

Type Material.—The female holotypein AMNH is labeled “Nan Siam [Nan,Thailand], January 13, Alice Mackie”; itis in good condition.

Distribution.—Known only from theholotype from Nan Province, Thailand.

Diagnosis, Comparative Comments,and Remarks.—Ceratina incertula wassynonymized by van der Vecht (1952)under C. collusor. However, van derVecht did not examine the type, butbased his opinion on Cockerell’s de-scription (1937) and concluded that C.incertula is a dark form of C. collusor.The type of C. incertula (examined by

NWand CDM) is not the same species asC. collusor, as shown by the presence ofpunctures along the parapsidal furrowand the area between the furrow and thelateral margin of scutum, and by theabsence of a protuberance on the outerside of the procoxa.

Ceratina incertula, however, is sosimilar to C. compacta that we are un-certain about its validity as a separatespecies. The punctation on the scutum ofC. incertula is consistent with some is-land populations (e.g., Luzon) of C.compacta in the Philippines, but differsfrom many including C. compacta fromThailand. In all Thai C. compacta, thescutum is heavily punctate, with almostno impunctate area; but in C. incertula,the impunctate area is obvious betweenthe anteromedian line and parapsidalfurrow. We retain C. incertula as a spe-cies separate from C. compacta for thetime being. More incertula-like speci-mens should be collected in Thailandto determine whether intergradationsexist that would indicate synonymy withC. compacta.

If C. incertula is found to be indeed C.compacta, the synonymy would indicatethat the species was in Thailand longbefore 1988; this would reject a majorcomponent in the hypothesis that thespecies is adventive in Thailand (Warrit2007b). Specimens considered to beC. compacta were not collected inThailand before 1988.

Ceratina lieftincki van der Vecht

Ceratina lieftincki van der Vecht, 1952:73–74 [male and female]

Type Material.—The male holotypeof C. lieftincki in RMNH (not seen) islabeled “Djasinga, 150 m., Tjibarangbang,15 Nov. 1936, resp. E. v. d. Vecht-B. andJ. v. d. Vecht”. The female paratype inthe same institution, is labeled “W. Java


Buitenzorg Mt. Tjiampea, 14 January1944, J. v. d. Vecht” [Java, Indonesia].The paratype is in fair condition; oneflagellum is missing, and one hind leg ismissing from femur to tarsus.

Thai Material Examined.—Bangkok:24 September 1969, Pensri (in Thai)(1♂; BKOK), 6 May 1973, P.S. Mes-senger (1♀; CAS); Chulalongkorn Uni-versity, 23 August 1999, Sonthaya (inThai) (1♀; BKOK). Chiang Mai: ChiangDoa District, 9–10 June 2003, 19°269N98°579E, N. Warrit (4♀, 2♂); ChiangMai University, 6 August 1988, Pornthep(in Thai) (1♀; CMAI); Mae Rim, 29January 1994, R.A. Beaver (1♀; LACM),26 January 1995, R.A. Beaver (1♀;LACM), 11 June 2003, 18°539N 98°519E,N. Warrit (1♂); Mae Wang, Don Pao, PaHuay Kho, 1–10 April 1997, 18°699N98°819E, Saowapa Sonthichai (2♀;LACM), 10–20 April 1997, 18°699N98°819E, Saowapa Sonthichai (2♀;LACM), 20–30 April 1997, 250 m,18°419N 98°489E, Saowapa Sonthichai(5♀; LACM). Chiang Rai: MeuangDistrict, 13 June 2003, N. Warrit (2♀),16 June 2003, 18°149N 99°299E, N.Warrit (4♀, 1♂). Chumpon: Chai YaDistrict, 14 July 2003, 09°199N 99°089E,N. Warrit (1♀); Lang Suan District, 15July 2003, 10°019N 99°039E, N. Warrit(4♀, 2♂); Meuang District, 25 June2003, 10°339N 99°079E, N. Warrit (4♀,2♂); Sapree District, 25 June 2003,10°349N 99°169E, N. Warrit (3♀, 2♂);Sawee District, 15 July 2003, 10°209N99°059E, N. Warrit (8♀). Kanchanaburi:110 air km. W. of Bangkok, 25–27 April1988, W.J. Pulawski (1♀; CAS); 10April 1989, W.J. Pulawski (1♀; CAS).Loei: 24 April 1989, W.J. Pulawski (1♀;CAS). Khon Kaen: Meuang District, 19June 2003, 16°269N 102°489E, N. Warrit(6♀, 6♂). Nakhon Nayok: NakhonNayok, 21 January 1993, S. Boongirdand C.D. Michener (2♀, 1♂; SEMC).

Nakhon Ratchasima: Burerum, 20 April1993, 163 m, Jarunya Thummanu (1♀;SEMC), 15 May 1993, 163 m, JarunyaThummanu (1♀; SEMC); Pak Chang,11 February 1993, S. Boongird and C.D. Michener (1♀; SEMC); SuranareeTechnological University, 20 June 2003,14°539N 102°009E, N. Warrit (8♀).Phanga: 14 June 2003, 08°389N 98°469E,N. Warrit (3♀, 5♂). Phattalung: 18 July2003, 07°229N 100°069E, N. Warrit(3♀). Phetburi: Khao Yoi District, 24June 2003, 13°159N 99°479E, N. Warrit(17♀). Phitsanulok: Bangraka District,17 June 2003, 16°449N 100°119E, N.Warrit (1♀). Prajinburi: Kabinburi,Bankhun Si, 5 December 1965, KolMongkolpanya (1♀; AMNH). PrachupKirikhan: Bangraka District, 26 June2003, 11°209N 99°299E, N. Warrit(4♀); Kui Buri District, 27 June 2003,12°059N 99°509E, N. Warrit (2♀, 1♂);Meuang District, 26 June 2003,11°409N 99°419E, N. Warrit (7♀); PranBuri District, 27 June 2003, 12°229N99°539E, N. Warrit (1♂). Rayong: KoSamet, 5–7 May 1989, W.J. Pulawski(2♂; CAS). Trang: Khao Chong NatureEducation Centre, 21– 24 July 1996,7°359N 99°469E, Snelling and Sonthichai(1♀; LACM); Sikoa District, 19 July2003, 07°319N 99°239E, N. Warrit(10♀, 4♂).

Distribution.—Ceratina lieftincki hasa wide range in Southeast Asia, from thePhilippines (Palawan Island) to In-donesia, Malay Peninsula, Thailand andCambodia.

Diagnosis and Comparative Com-ments.—Ceratina lieftincki is similarto C. compacta, but it can be distin-guished from the latter by the absenceof yellow marking on the base of theantennal scape and by the sparsely punc-tate (interspaces broader than puncturediameters) posterolateral area of the mes-episternum near the mesocoxal base.


Ceratina nigrolateralis Cockerell

Ceratina philippinensis nigrolateralis Cockerell,1916: 305 [female];Ceratina nigrolateralis,van der Vecht, 1952: 77–81 [recognized asa species; redescription of female and male;key to subspecies].

Ceratina acuticauda Cockerell, 1919b: 175[male].

Ceratina incerta Cockerell, 1919a: 247 [femaleand male]; Cockerell, 1920a: 624 [men-tioned in key]; Cockerell, 1929: 150 [newrecord from Thailand (Siam)].

Ceratina corbetti Cockerell, 1929: 151 [fe-male, male].

Type Material.—The female holotypein BMNH is labeled “Ceratina philip-pinensis nigrolateralis”, “T.D.A. CockerellB.M. 1934-527”, “3887”, “P. PrincesaPalawan Baker” [Palawan, Island, Philip-pines], “Type # 17 b 204”. The type is ingood condition; it has one row of puncturesalong the parapsidal furrow and yellowmarking on the base of the antennal scape.

The male holotype of C. acuticauda inUSNM is labeled “Type No. 20705 U.S.N.M.”, “Buitenzorg [= Bogor] Java 4.09”, and “Bryant & Palmer Coll” [Java,Indonesia]. The type is in good condi-tion. The holotype of Ceratina acuti-cauda is a male of C. nigrolateraliswithout yellow on the base of the scape.

The female holotype of C. incerta inBMNH is labeled “T.D.A. Cockerell B.M. 1934–527”, “Singapore Coll. Baker”,and “Type # 17 b 236”. The type is ingood condition. Ceratina incerta isa dark form of C. nigrolateralis with themesotibiae and metatibiae black.

The female holotype of C. corbetti inUSNM is labeled “Kuala Lumpur 7 ms1928 Feb (Cockerell)” [Malaysia] and“Type No. 58143 U.S.N.M.”. The type isin good condition. Van der Vecht (1952)synonymized C. corbetti under C. ni-grolateralis, but retained C. acuticaudaand C. corbetti as different subspecies.

We have examined the type of C. cor-betti and decided that it is a female of C.acuticauda [= C. nigrolateralis] withreduced yellow markings on the meso-tibiae and metatibiae. We do not recog-nize any of these morphs as subspecies.

Thai Material Examined.—Bangkok:17 April 1973, on flowers of Ziziphus, P.S. Messenger (1♀; CAS); 19 May 1973,on flowers, P.S.Messenger (8♀, 1♂; CAS),6 June 1973, on flowers, P.S. Messenger(2♀; CAS). Chantaburi: ; 2 km. S. ofChantaburi, 19–21 June 1969, 12 369N102 079E, J.J.S. Burton (1♀; BPBM).Chiang Mai: Amphur Mae Wang,Thanbon Donpao, Pa Huay Kho, 20–30April 1997, 18° 41.549N 98° 48.4179E,250 m., Malaise trap in field at edge ofdeciduous forest, Saowapa Sonthichai(1♀; LACM); Chiang Doa District, 9–10June 2003, 19°269N 98°579E, N. Warrit(14♀, 1♂); Mae Rim District, 11 June2003, 18°539N 98°519E, N. Warrit (14♀,7♂). Chiang Rai: Meuang District, 12June 2003, 20°059N 99°519E, N. Warrit(2♀); 13 June 2003, N. Warrit (22♀).Chumpon: Chai Ya District, 14 July 2003,09°199N 99°089E, N. Warrit (1♀, 5♂);Lang Suan District, 15 July 2003,10°019N 99°039E, N. Warrit (12♀, 5♂);Meuang District, 25 June 2003, 10°339N99°079E, N. Warrit (8♀, 1♂); SapreeDistrict, 25 June 2003, 10°349N 99°169E,N. Warrit (5♀); Sawee District, 15 July2003, 10°209N 99°059E, N. Warrit(5♀, 2♂). Kanchanaburi: 110 air km.W. of Bangkok, 25–27 April 1988, W.J.Pulawski (2♀; CAS). Khon Kaen:Meuang District, 19 June 2003, 16°269N102°489E, N. Warrit (1♀). NakhonNayok: Nakhon Nayok, 21 January 1993,S. Boongird and C.D. Michener (2♀, 1♂;SEMC). Nakhon Ratchasima: Tha Chang,10 February 1993, ex. Callistemon, S.Boongird andC.D.Michener (1♀; SEMC).Phanga: 14 June 2003, 08°389N 98°469E,N. Warrit (2♀). Phetburi: Khao Yoi


District, 24 June 2003, 13°159N99°479E, N. Warrit (5♀). Phuket: PhuketIsland, 29 January 1993, S. Boongirdand C.D.Michener (2♀; SEMC). Saraburi:August, Sakda (in Thai) (1♀; BKOK).Ranong: Kuraburi District, 16 July 2003,09°199N 98°259E, N. Warrit (1♀, 11♂);Meuang District, 16 July 2003, 09°519N98°379E, N. Warrit (2♀). Surat Thani:Bang-O 25 km. SW. of Surat Thani, 30January 1993, S. Boongird and C.D.Michener (1♀; SEMC). Trang: KhaoChong Nature Education Center, 21–24July 1996, 07 °359N 99°469E, Snellingand Sonthichai (3♀; LACM); SikoaDistrict, 19 July 2003, 07°319N 99°239E,N. Warrit (9♀, 1♂).

Distribution.—Ceratina nigrolater-alis is widely distributed throughoutSoutheast Asia—Laos, Thailand, Viet-nam, Malaysia, Singapore, Indonesia,and Palawan Island, Philippines.

Diagnosis and Comparative Com-ments.—Ceratina nigrolateralis can bedistinguished from other Ceratinidia inthe compacta species group by the ab-sence of punctures along the parapsidalfurrows and in the area between thefurrow and the lateral margin of the scu-tum, and in the presence of a protuberanceon the outer side of the procoxa.

Ceratina nigrolateralis is similar to C.collusor in many respects, but can bedifferentiated by the relatively large sizeand the absence of yellow marking onthe base of the scape (present in Philip-pine population). Van der Vecht (1952)segregated four subspecies in C. nigro-lateralis: C. nigrolateralis nigrolateralis(Palawan Island, Philippines), C.nigrolateralis acuticauda (Thailand,Laos, Vietnam, Malay Peninsula, andIndonesia), C. nigrolateralis incerta(Singapore and Sumatra), and C. ni-grolateralis corbetti (Indochina, MalayPeninsula, and Indonesia). However, weconsider C. nigrolateralis acuticauda and

C. nigrolateralis corbetti to be a singlemorph, and do not recognize the others assubspecies.

Ceratina sutepensis Cockerell,new status(Figs. 8, 9)

Ceratina lepida var. sutepensis Cockerell, 1929:150 [female]; van der Vecht, 1952: 54[notes on type].

Ceratina lepida var. sublepida Cockerell,1929: 150 [female]; van der Vecht, 1952:54 [notes on type] new synonym.

Type Materials.—A female type of C.lepida var. sutepensis in BMNH, is la-beled “Ceratina lepida var. sutepensisCockerell, 1929.”, “B.M. Type Hym. 14b 220”, and “Doi Suthep, Siam”. Theyellow maculation of this type is muchreduced, especially on the pronotum,pronotal lobe, mesoscutum, and mesepis-ternum. Since neither Cockerell nor sub-sequent authors designated the “type”, wetherefore designate as lectotype the femaletype of C. lepida var. sutepensis inBMNH, and add a red label “Ceratinalepida var. sutepensis Cockerell, Lecto-type, N. Warrit, 2008”.

A female type of C. lepida var. sub-lepida is similar to the type of C. lepidavar. sutepensis in punctation (note that ofclypeus and paraocular area); however,the metasoma of the C. lepida var. sub-lepida type is missing. The type of C.lepida var. sublepida is labeled “Cera-tina lepida var. sublepida Ckll.”, “DoiSutep, Siam”, and “B.M. Type Hym. 14b 222”. We designate this specimen aslectotype, and added a red label “Cera-tina lepida var. sublepida Cockerell,Lectotype, N. Warrit, 2008”.

The types of C. lepida var. sutepensisand var. sublepida were both collectedfrom the same area, Doi Suthep. Thetwo names, originally “varieties” ofC. lepida, were published simultaneously.


As first revisers, we have selectedsutepensis as the valid name for thespecies. The difference between the twois only in the extent of yellow macula-tion, especially on the pronotum andscutellum.

Thai Material Examined.—ChiangMai, Ahn Kang, W. of Fang, 6 February1993, S. Boongird, C. D. Michener, andS. Malaipan (1♀; SEMC); Chiang Dao, 5–11 April 1958, T. C. Maa (1♀; BPBM);Doi Suthep, 22 April 1988, W. J. Pulawski(1♀; CAS); Poi Suthee, 21 August 1990,G. W. Otis (1♀, 2♂; SEMC).

Distribution.—Ceratina sutepensis isknown only from the mountainous areaof Chiang Mai Province, Thailand.

Diagnosis and Comparative Com-ments.—The female of C. sutepensis is

similar to that of C. bryanti, but the yel-low markings are much more reduced,especially on the pronotum, pronotallobe, scutum, scutellum, and terga. Thefacial punctation ofC. sutepensis, notablyin the female, is unique among Ceratini-dia except in the bryanti species group.The pattern of punctation on the para-ocular area is with fine to coarse punc-tures clumped together on the medianpart, but with only a few punctures on thelower and upper parts. The clypeus hasa conspicuous median longitudinal im-pressed area, but it is not as pronouncedas in C. hieroglyphica.

Themale ofC. sutepensis is also similarto that ofC. bryanti but with more reducedmaculation. Markings on the followingstructures are absent in C. sutepensis,

Figs. 8–11. Ceratina (Ceratinidia) sutepensis Cockerell (left side) and C. bryanti Cockerell (rightside), lateral views and faces.


while present in C. bryanti: pronotal lobe(reduced), transverse marking on prono-tum, scutellum, metatibia, and terga. Theapex of T7 of both species is pointed withangulated margin, but in C. bryanti theapex is more produced with a conspicu-ous emargination.

ACKNOWLEDGMENTS

We thank Tipwan Suppasat, MarutFuangarworn, and Asalek Ratanawanneefor their help in the field. Lynn S. Villafuerteand Joseph A. David provided helpfulassistance in specimen recording. Thephotographic equipment was providedby the Smithsonian Institution and theSystematic Entomology Laboratory (SEL,USDA) through Mathew L. Buffingtonand David G. Furth. For organization ofthe illustrations we are entirely indebted toIsmael A. Hinojosa-Diaz.

We thank the following people andtheir institutions for providing specimensfor study: Cornelius van Achterberg, JohnS. Ascher, George R. Else, Michael S.Engel, Zachary H. Falin, David G. Furth,Tino Gonsalves, Maureen Melo, WojciechJ. Pulawski, Jerome G. Rozen Jr., G. AllanSamuelson, the late Roy R. Snelling,Prachaval Sukhumalanand, and Yan-ruWu. The research by CL and NW waslargely supported from the BiodiversityResearch Training Program (BRT-149006), Thailand, and that by NW, fromthe Hungerford Fund and the RaymondBeamer Fellowship of the University ofKansas Entomology Program, USA, andthe Thailand Research Fund (TR♀MRG5380139) andGrants forDevelopmentof New Faculty Staff, Chulalongkorn Uni-versity, Thailand.

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