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JHR 42: 63-92 (2015) JOURNAL OF *0eerriewed opevarcets oral
Pie ees (4) Hymenoptera

http://jhr.pensoft.net The insertional Society of ymenoptersts. RESEARCH

Revision of the Crematogaster ranavalonae-group
in Asia, with description of two new species
(Hymenoptera, Formicidae)

Shingo Hosoishi!

| Institute of Tropical Agriculture, Kyushu University, Fukuoka, 812-8581 Japan

Corresponding author: Shingo Hosoishi (hosoishi@gmail.com)

Academic editor: W. Pulawski | Received 16 October 2014 | Accepted 4 February 2015 | Published 18 March 2015
Attp.//zoobank.org/7674B232-8BED-4BA0-B4FE-D22A9D2A1C45

Citation: Hosoishi S (2015) Revision of the Crematogaster ranavalonae-group in Asia, with description of two new species
(Hymenoptera, Formicidae). Journal of Hymenoptera Research 42: 63-92. doi: 10.3897/JHR.42.8758

Abstract

The Asian members of the Crematogaster ranavalonae-group are revised and twelve species, including two
new species, C. hashimi sp. n. and C. imperfecta sp. n. are recognized. The members are distinguished
from the other Asian Crematogaster in having smooth, shiny bodies with short appressed setae on the sur-
face. Crematogaster sikkimensis Forel, 1904 is raised to the species level, and the following new synonyms
are established: C. aberrans Forel, 1892 = C. aberrans assmuthi Forel, 1913, syn. n. = C. aberrans inglebyi
Forel, 1902, syn. n. = C. sovor Forel, 1902, syn. n.; C. ebenina Forel, 1902 = C. ebenina corax Forel, 1902,

syn. n. A key to the species based on the worker caste is provided.

Keywords
Asia, Crematogaster ranavalonae-group, Formicidae, new species, Oxygyne, taxonomy, ventrolateral

katepisternal ridge

Introduction

The hyperdiverse ant genus Crematogaster had been subdivided into sixteen subgenera
(Emery 1922; Wheeler 1922). Recently a molecular phylogenetic analysis has sug-
gested that most of these subgenera are non-monophyletic, and that the subgenus
Oxygyne is monophyletic (Blaimer 2012c). Subsequently Blaimer (2012b) established

Copyright Shingo Hosoishi. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

64 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

the Crematogaster ranavalonae-group to accommodate the former subgenus Oxygyne in
her subgeneric revision of this genus.

Members of the Crematogaster ranavalonae-group have been differentiated based
on characters found in the worker and queen castes (Blaimer 2012a, 2012b). Although
workers of the Asian members also posses the taxonomic characters identified by her,
the propodeal spines in some of the Asian species are not well developed, appearing
instead as small tubercles. However, I have identified a unique character among the
members of the Asian C. ranavalonae-group; that is, while the ridge separating the la-
teral and ventral portions of the mesopleuron is distinct in most Crematogaster species,
it is not distinct in the Asian members of the C. ranavalonae-group. Furthermore, the
queen caste of some members of the species group has falcate mandibles, suggesting the
occurrence of temporal social parasitism in those species (Forel 1910; Santschi 1934;
Holldobler and Wilson 1990; Blaimer 2012a). However, among the Asian fauna, falca-
te mandibles are found only in the queen of C. augusti and are unknown in other species
due to the rarity with which they are encountered in the field. In this study, I do not
treate the queen caste because it is not represented in my collections.

The Crematogaster ranavalonae-group consists of twenty-two species, including ele-
ven species from Africa and Madagascar, ten from Asia, and one from New Guinea (Blai-
mer 2012b). Among the Asian fauna (ten species and four subspecies), four species and
three subspecies have been described from India. It is considered that India is the center
of diversity in this species-group, but this may be an overestimate attributable to the rari-
ty of this species group in the field. This paper provides a revision of the Asian members
of C. ranavalonae-group, based on the morphological characters of the worker caste.

Materials and methods

Sources of material
Specimens were examined and/or deposited in the collections listed below. Codes for
public institutions mainly follow those in Brandao (2000). Nest series samples, most

of which were recently collected, are represented as colony codes, e.g., “SH12-Tha-01.”

BMNH The Natural History Museum, London, U. K.

CASC California Academy of Sciences, San Francisco, CA, USA

FRIM Forest Research Institute Malaysia, Kepong, 52109 Kuala Lumpur,
Malaysia.

IEGG Istituto di Entomologia “Guido Grandi”, Bologna, Italy.

KUM Kyushu University, Fukuoka, Japan.

MCSN Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy.

MCZC Museum of Comparative Zoology, Harvard University, Cambridge,
MA, USA.

MHNG Musée d’Histoire Naturelle, Geneva, Switzerland.

Revision of the Crematogaster ranavalonae-group in Asia... 65

MBBJ Museum Zoologicum Bogoriense, Cibinong, Java, Indonesia.

NHMB Naturhistorisches Museum, Basel, Switzerland.

NHMW Naturhistorisches Museum Wien, Wien, Austria.

SKYC Sk. Yamane Collection, Kagoshima University, Japan.

THNHM Thailand Natural History Museum, Technopolis, Khlong Luang,
Pathum Thani, Thailand.

Methods

Most observations were made on an Olympus SZX12 stereomicroscope. The scanning
electron micrographs (SEM) were prepared with a JSM-5G600LYV scanning electron
microscope. Color images were taken using a Canon EOS 50D with a Canon MP-E
65 mm 1—5x macro lens, then processed using Combine ZM.

The relative position of propodeal spiracles was observed in lateral view, with the
mesosoma carefully tilted to the position with the true maximum of Weber’s length.

Measurements and indices

Measurements were made under an Olympus SZX12 stereomicroscope using microm-
eters. All measurements are expressed in millimeters, recorded to the second decimal
place. The measurements for petiole and postpetiole follow Longino (2003).

Head Width (HW): Maximum width of head in full-face view, excluding the eyes
(Fig. 1).

Head Length (HL): Perpendicular distance from vertex margin to line tangent to
anteriormost projections of clypeus in full-face view (Fig. 1).

Cephalic Index (CI): HW/HL x 100.

Scape Length (SL): Length of the first antennal segment, excluding the neck and
basal condyle (Fig. 2).

Scape Index (SI): SL/HW x 100.

Eye Length (EL): Maximum length of the compound eye (Fig. 1).

Pronotal Width (PW): Maximum width of the pronotum in dorsal view (Fig. 3).

Weber’s Length of the mesosoma (WL): Diagonal length, measured in lateral view
from the anterior margin of the pronotum (excluding the collar) to the posterior ex-
tremity of the propodeal lobe (Fig. 4).

Propodeal Spine Length (PSL): measured from tip of propodeal spine to closest
point on outer rim of propodeal spiracle (Fig. 4).

Petiole Length (PtL): Length of the petiole in lateral view (Fig. 5) (see Longino
2003 ;<fie.2),

Petiole Width (PtW): Maximum width of petiole in dorsal view (Fig. 6).

Petiole Height (PtH): Height of the petiole in lateral view (Fig. 5) (see Longino
2003, fig. 2).

66 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

5 6 a PpL

Figures |-—6. Measurements of workers. | Head width, head length and eye length 2 Scape length
3 Pronotal width 4 Weber’s length and propodeal spine length 5 Petiole length and petiole height 6 Peti-
ole width, postpetiole length and postpetiole width.

Postpetiole Length (PpL): Length of the postpetiole in dorsal view (Fig. 6) (see
Longino 2003, fig. 2).

Postpetiole Width (PpW): Maximum width of postpetiole in dorsal view, excluding
the helcium (Fig. 6).

Revision of the Crematogaster ranavalonae-group in Asia... 67

Figure 7. Ventrolateral katepisternal ridge from ventral portion in mesosoma (legs removed). Arrow

indicates the ridge. A Crematogaster hashimi B Crematogaster physothorax.

Petiole Height Index (PtHI): PtH/PtL x 100.

Petiole Width Index (PtWI): PtW/PtL x 100.

Postpetiole Width Index (PpWI): PpW/PpL x 100.

Waist Index (WI): PpW/PtW x 100.

An important character in the Asian Crematogaster ranavalonae-group is explained below.

Ventrolateral katepisternal ridge (Fig. 7). The ridge separates the mesopleuron lat-
eral surface from ventral surface. Most Asian Crematogaster species have a well-defined
ridge separating the two surfaces (Fig. 7B), but in some species the ridge is absent or ves-
tigial (Fig. 7A). The ridge is visible in pinned specimens from lateral or ventrolateral view.

Asian Crematogaster ranavalonae-group

The Asian members of the Crematogaster ranavalonae-group not only have the mor-
phological features diagnosed by Blaimer (2012a, 2012b), but also show the additional
features below.

(i) Pronotum steeply raised in lateral view.

(ii) Ventrolateral katepisternal ridge indistinct, but weakly developed anteriorly in
some species.

(iii) Integument essentially smooth and shiny.

(iv) Erect pilosity almost absent. Some erect setae are developed on the clypeus or
dorsal surface of petiole and postpetiole, but absent on the dorsum of head, mes-
osoma and fourth to seventh abdominal tergites.

(v) Dorsum of head, mesosoma and fourth abdominal tergite with short and ap-
pressed setae.

This species group is easily defined from other Asian Crematogaster in having a steeply
raised pronotum, smooth and shiny body surface, and short and appressed body setae.

68

Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

Synonymic list of the Asian Crematogaster ranavalonae-group

aberrans Forel, 1892.

= aberrans assmuthi Forel, 1913, syn. n.
= aberrans inglebyi Forel, 1902, syn. n.
= soror Forel, 1902, syn. n.

augusti Emery, 1895.
butteli Forel, 1913.
daisyi Forel, 1901.
dalyi Forel, 1902.
ebenina Forel, 1902.

= ebenina corax Forel, 1902, syn. n.

hashimi sp. n.

imperfecta sp. n.

pia Forel, 1911.

sitkkimensis Forel, 1904. stat. n.
tumidula Emery, 1900.
vandermeermohri Menozzi, 1930

Key to species based on the worker caste (not included are C. augusti, known only from

the queen caste, and C. vandermeermohri whose type specimens were not examined)

1

Scape with sparse setae (Fig. 8). Propodeal spiracles as small as mesothoracic
Spiracleg SUP IG eT On kadai cath acl Nauman tiaailaelgaweors eteaRoxaien staat nat tastonisusds aberrans
Scape with abundant setae (Fig. 9). Propodeal spiracles large; diameter ca. 2

%.as latee as-mesorhbracic SpiraclesA Fig. FL): |. ceuesnsecrosstvonerstnenedtereandecinante 2
Postpetiole markedly bilobed laterally, more than 2 x as broad as long (PpWI
aD E YG We Ui Sar sk MOR eR she vit tin Magers Hed py tera Pes) ae Be 3
Postpetiole moderately bilobed laterally, generally ca. 1.2 to 2 x as broad as
Lone CP PED F220 CR eSal Det A Mie, a tesenerath oeniaisentea net tlacetniss igh ieeeirenonedabi 4
Propodeal spines short; their length equal to or shorter than diameter of pro-
podedlspitacles iG. 1) tic. ddanwesk teat ebnannutdedtnatasthicnsesiteat ea vcebuiasaeu. dalyi
Propodeal spines long; distinctly longer than diameter of propodeal spiracles
PISS). ccouve cocenerscurvevccuerecucoretvcenecctermecneerccniseyereutincxcoueaed sikkimensis
Propodeal spines not developed, reduced to small tubercles (Figs 17, 18) ...5
Rropodeall spines;develo ped GRie N rssucrastveestensieovbautuaseys wees) enon sereseethan 6
Propodeal spiracles circular (Fig. 17). Petiole about as broad as or broader
thandong MPEWA DG—109) CPI ELOY a cers selcrasnsenonsntnoop oadsopeunnteh vas ceed hashimi
Propodeal spiracles elliptical (Fig. 18). Petiole longer than broad (PtWI 88-
D2) UE SI tt ates axa Raa nel aoe eealnn casi tno Paladin on tose tea dean bee snal imperfecta

Anterolateral portion of petiole developed laterally and spiracles on petiole
located on lower position and directed ventrally (Fig. 21)... daisyi

Revision of the Crematogaster ranavalonae-group in Asia... 69

0.2 mm
Sn ieenitieiimateaienianm

Figures 8-25. Worker characters of Asian Crematogaster ranavalonae-group. 8 left scape (C. aberrans)
9 left scape (C. daisyi) 10 propodeal spiracle (C. aberrans) 11 propodeal spiracle (C. daisyi) 12 petiole
and postpetiole (C. dalyi) 13 petiole and postpetiole (C. ebenina) 14 petiole and postpetiole (C. pia)
15 propodeal spine (C. dalyi) 16 propodeal spine (C. sikkimensis) 17 propodeal spiracle and reduced
propodeal spine (C. Aashimi) 18 propodeal spiracle and reduced propodeal spine (C. imperfecta)
19 petiole and postpetiole (C. Aashimi) 20 petiole and postpetiole (C. imperfecta) 21 spiracle on petiole
(C. daisyi) 22 spiracle on petiole (C. ebenina) 23 petiole and postpetiole (C. butteli) 24 propodeal spine
(C. tumidula) 25 propodeal spine (C. ebenina).

70 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

= Anterolateral portion of petiole developed laterally, but spiracles on petiole
located on middle portion and directed laterally (Fig. 22)... eee eeeeeeeeees 7
7 Anterolateral corners of petiole angulate, anterior face developed (Fig. 23) . butteli
Anterolateral corners of petiole rounded, anterior face sinuate (Fig. 13)......8
8 Propodeal spines short and stout; as long as diameter of propodeal spiracles

CE) te reais cecil: Eade ope ant te eeuee Maan cBarrsunsoue ene Me ecloeee wotenstourene tumidula

9 Anterolateral corners of petiole undeveloped, tapering anteriorly (Fig. 13).
Postpetiole generally as wide as petiole in dorsal view (WI 96-100) ... ebenina
— Anterolateral corners of petiole developed (Fig. 14). Postpetiole wider than
petiolevim dorsaltsri cys Wil 03s S) iis iaceed ts sce eereen ted nob estes dedaabeteeses pia

Species accounts

Crematogaster aberrans Forel

Fig. 26

Crematogaster aberrans Forel, 1892: 532-534. Type locality: India, Thana. Combination
in C. (Oxygyne) by Forel, 1901: 375; in C. (Crematogaster) by Blaimer, 2012b: 55.

Crematogaster (Oxygyne) aberrans var. assmuthi Forel, 1913: 662. Type locality: India,
Bombay. syn. n.

Crematogaster (Oxygyne) aberrans var. inglebyi Forel, 1902: 201. Type locality: India,
Travancore. syn. n.

Crematogaster (Oxygyne) soror Forel, 1902: 200. Type locality: India, Poona and
Bombay. Combination in C. (Crematogaster) by Blaimer, 2012b: 55. syn. n.

Type material examined. Crematogaster aberrans: lectotype worker (top specimen of
three on one pin) by present designation and five paralectotype workers from India,
Thana (MHNG, examined); one syntype worker from India, Thana (NHMB, exam-
ined). Crematogaster (Oxygyne) aberrans var. assmuthi: one syntype worker from India,
Bombay (M. Assmuth) (MHNG, examined). Crematogaster (Oxygyne) aberrans vat. in-
glebyi: three syntype worker from India, Travancore (Ferguson and Ingleby) (MHNG,
examined). Crematogaster (Oxygyne) soror: five syntype workers from India, Poona
(Wroughton) and Bombay (Rothney) (MHNG, examined); one syntype worker from
India, Bombay (Rothney) (NHMB, examined).

Non-type material examined. THAILAND: 8 workers, Doi Chiang Dao (500-
600m alt.), nr Chiang Mai, N. Thailand. 2.iv.2005 (TH05-SKY-22) (Sk. Yamane); 5
workers, Ngao, Lampang, 17.ii.1991 (Vo collector’s name); 4 workers, Maegar, Phayao,
30.i1i1.1991 (No collector’s name).

Measurements and indices (workers, n = 10). HW 0.82-0.96; HL 0.78-0.93; CI
102-109; SL 0.64-0.74; SI 75-82; EL 0.18-0.21; PW 0.50-0.58; WL 0.90-1.05; PSL

Revision of the Crematogaster ranavalonae-group in Asia... Fl

Figure 26. Crematogaster aberrans [Doi Chiang Dao, nr. Chiang Mai, N. Thaiand]. A Lateral view of
body B Full face view C Dorsal view of petiole and postpetiole.

0.09-0.12; PtL 0.25-0.30; PtW 0.25-0.32; PtH 0.17-0.20; PpL 0.15-0.19; PpW 0.25-
0.33; PtHI 61-76; PtWI 97-116; PpWI 167-213; WI 100-114.

General description of worker. Head appearing rounded in front view. Mandible
indistinctly striate, with four teeth, apical and subapical teeth large, basal two teeth
smaller. Scape exceeding posterior corner of head, with sparse appressed setae, each of
which is about as long as scape diameter. Compound eye slightly projecting beyond
lateral margin of head in full face view.

Mesonotum highly convex in lateral view. Ventrolateral katepisternal ridge in-
distinct. Propodeal spine short and stout; the length variable, but generally as long as
propodeal spiracle, dorsum not higher than anterior propodeum in lateral view. Pro-

72 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

podeal spiracle as large as mesothoracic spiracle pit, situated apart from the propodeal
declivity in lateral view, directed laterally.

In dorsal view, shape of petiole scoop with convex side, as long as broad. Petiole
with anterolateral corner angulate. Petiolar spiracle big, as wide as half of propodeal
spiracle in diameter, directed laterally. Postpetiole bilobed, with slight longitudinal
median sulcus. Postpetiole slightly wider than petiole in dorsal view.

Integument essentially smooth and shiny. Clypeus generally smooth and shiny,
but weakly striated with faint rugulae. Malar region with feeble longitudinal rugu-
lae. Dorsal surface of promesonotum weakly punctuated. Lateral surface of pronotum
shiny, but weakly punctuated. Mesopleuron weakly punctuated. Dorsal surface of pro-
podeum smooth and shiny. Lateral propodeum smooth and shiny.

Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short
and appressed, sparse setae. Clypeus without pair of longer setae on anteriormost por-
tion. Anterior clypeal margin with one single longer seta and one pair of longer setae
on median portion, mixed with some shorter setae on side. One to three pairs of short,
erect setae on pronotal shoulder. Posterolateral tubercle with two pairs of erect setae.
Ventral surface of petiole with longer appressed setae. Postpetiole with two pairs of
longer setae posteriorly. Fourth abdominal tergite with appressed setae.

Body color reddish brown.

Distribution. This species is distributed in India and Thailand.

Comments. This species is unique among Asian Crematogaster ranavalonae-group
in having the scape with sparse setae and smaller propodeal spiracles, situated apart
from the propodeal declivity.

The syntype workers of C. aberrans assmuthi match well with syntype workers of C.
aberrans. | treated C. aberrans assmuthi as a junior synonym of C. aberrans. The syntype
workers of C. aberrans inglebyi match well with syntype workers of C. aberrans, but
with only the following small difference: the former has right brown colored body (cf.
Forel 1902). I concluded that C. aberrans inglebyi is a junior synonym of C. aberrans.
The syntype workers of C. soror match well with syntype workers of C. aberrans. In
Forel’s original description (Forel 1902), he mentioned that the head is subquadratic.
However, the posterior margin of head is not angulate, appearing rounded in front
view. Thus I here synonymize C. soror with C. aberrans.

Crematogaster augusti Emery

Crematogaster augusti Emery, 1895: 31, pl. 2, figs. 19, 20. Type locality: Indonesia,
Marang, Sumatra. Combination in C. (Oxygyne) by Emery, 1922: 157; in C. (Cre-
matogaster) by Blaimer, 2012b: 55.

Type material examined. Crematogaster augusti: one syntype queen from Indonesia,

Marang, Sumatra (MCSN, examined).

Revision of the Crematogaster ranavalonae-group in Asia... 73

Comments. Emery’s original description was based on a single queen. Our collec-
tions are limited to worker specimens, but two unique features are found in the type.
First, the propodeal spines are not developed, as they are in the queen of C. agnetis Forel
from Madagascar (Blaimer 2012a). Second, the body surface appearing smooth and
shiny. The taxonomic status will remain uncertain until nest series become available.

Crematogaster butteli Forel
Fig.27

Crematogaster (Oxygyne) butteli Forel, 1913: 78. Type locality: Indonesia, Soengei
Bamban, Sumatra. Combination in C. (Crematogaster) by Blaimer, 2012b: 55.

Type material examined. Crematogaster (Oxygyne) butteli: lectotype worker (top speci-
men of two on one pin) by present designation and four paralectotype workers from
Indonesia, Soengei Bamban, Sumatra (MHNG, examined); one syntype worker from
Indonesia, Soengei Bamban, Sumatra (NHMB, examined).

Measurements and indices (type workers, n = 5). HW 0.80-0.85; HL 0.76-0.80;
CI 103-107; SL 0.64-0.67; SI 79-84; EL 0.16-0.17; PW 0.48-0.52; WL 0.91-0.96;
PSL 0.12-0.14; PtL 0.21-0.22; PtW 0.22-0.24; PtH 0.15-0.16; PpL 0.14-0.15; PpW
0.24-0.25; PtHI 68-76; PtWI 95-114; PpWI 167-179; WI 100-119.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two
teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of
which is as long as width of scape. Compound eye large and slightly projecting beyond
lateral margin of head in full face view.

Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine long and
stout, longer than spiracle diameter, directed upward, dorsum higher than anterior
propodeum in lateral view. Propodeal spiracle large, situated on lateral surface of pro-
podeum and close to propodeal declivity in lateral view.

In dorsal view, shape of petiole scoop with tapering side posteriorly, longer than
broad. Petiole with anterolateral corner angulate. Petiolar spiracle big, as wide as half
of propodeal spiracle in size. Postpetiole bilobed but without longitudinal median
sulcus. Postpetiole as wide as petiole in dorsal view.

Integument essentially smooth and shiny. Clypeus smooth and shiny without ru-
gulae. Malar region generally smooth, with feeble rugulae. Dorsal surface of prome-
sonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Meso-
pleuron shiny, but with longitudinal rugulae. Dorsal surface of propodeum smooth
and shiny. Lateral propodeum shiny, but with longitudinal rugulae.

Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with sparse,
short and appressed setae. Clypeus with one pair of longer setae on anteriormost por-
tion, directed medially. Anterior clypeal margin with two pairs of longer setae on

74 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

Figure 27. Crematogaster butteli [Soengei, Bamban, Sumatra, Indonesia]. A Lateral view of body B Full

face view C Dorsal view of petiole and postpetiole.

median portion, mixed with some shorter setae on sides. No erect setae on pronotal
shoulder. Posterolateral tubercle with some appressed setae. Ventral surface of petiole
with appressed setae. Postpetiole with one pair of longer erect setae posteriorly. Fourth
abdominal tergite with appressed setae.

Body color reddish brown.

Distribution. This species is known only from the type locality in Sumatra.

Comments. This species is similar to C. tumidula but can be distiguished in having
angulate anterolateral corners of the petiole .

Revision of the Crematogaster ranavalonae-group in Asia... 75

Figure 28. Crematogaster daisyi [Tasek Merimbun, Brunei]. A Lateral view of body B Full face view

C Dorsal view of petiole and postpetiole.

Crematogaster daisyi Forel
Fig. 28

Crematogaster (Oxygyne) daisyi Forel, 1901: 376. Type locality: Malaysia, Sarawak,
Borneo. Combination in C. (Crematogaster) by Blaimer, 2012b: 55.

Type material examined. Crematogaster (Oxygyne) daisyi: lectotype worker (bottom
specimen of three on one pin) by present designation and five paralectotype work-
ers from Malaysia, Sarawak, Borneo (Haviland) (MHNG, examined); three syntype
workers from Malaysia, Sarawak (NHMW, examined); three syntype workers from
Malaysia, Sarawak (Haviland) (NHMB, examined).

76 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

Non-type material examined. MALAYSIA: 1 worker, Lambir National Park,
Miri, Sarawak, Borneo, 2.i1.1998, (S&. Yamane); BRUNEI: 1 worker, Tasek Merim-
bun, 11.ii.1999 (Eg99-BOR-011) (K. Eguchi); 1 worker, Tasek Merimbun, 17.ii.1999
(Eg99-BOR-155) (K. Eguchi).

Measurements and indices. (workers, n = 6). HW 0.79-0.88; HL 0.74-0.83; CI
101-107; SL 0.68-0.76; SI 86-90; EL 0.17-0.18; PW 0.49-0.54; WL 0.91-0.97; PSL
0.11-0.15; PtL 0.22-0.24; PtW 0.22-0.26; PtH 0.13-0.15; PpL 0.15-0.17; PpW 0.27-
0.33; PtHI 59-68; PtWI 100-130; PpWI 169-200; WI 107-136.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two
teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of
which is as long as width of scape. Compound eye large and slightly projecting beyond
lateral margin of head in full face view.

Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine long and
stout; length greater than spiracle, directed upward, dorsum higher than anterior pro-
podeum in lateral view. Propodeal spiracle large, touching to propodeal declivity in
lateral view, directed laterally.

In dorsal view, shape of petiole scoop with convex side, longer than broad. Petiole
with subparallel side on anterior half in dorsal view. Petiolar spiracle large, as large as
half of propodeal spiracle, directed downward. Postpetiole bilobed with longitudinal
median sulcus. Postpetiole wider than petiole in dorsal view.

Integument essentially smooth and shiny. Clypeus smooth and shiny without rugulae.
Malar regions generally smooth. Dorsal surface of promesonotum smooth and shiny.
Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral propodeum
generally shiny, but with feable rugulae. Dorsal surface of propodeum smooth and shiny.

Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short
and appressed setae sparsely. Clypeus with one pair of longer setae on anteriormost
portion, directed medially. Anterior clypeal margin with two pairs of longer setae,
mixed with some shorter setae on sides. No erect setae on pronotal shoulder. Poste-
rolateral tubercle with some decumbent to appressed shorter setae. Ventral surface of
petiole with appressed setae. Postpetiole with some shorter setae posteriorly. Fourth
abdominal tergite with appressed setae.

Body color reddish brown.

Distribution. This species is only known from Malaysia (Borneo).

Comments. ‘This species is similar to C. sikkimensis, but differs in having the spira-
cles on the petiole located on the lower position in lateral view.

Crematogaster dalyi Forel
Fig. 29

Crematogaster (Oxygyne) dalyi Forel, 1902: 201. Type locality: India, Coonoor. Com-
bination in C. (Crematogaster) by Blaimer, 2012b: 55.

Revision of the Crematogaster ranavalonae-group in Asia... 77

Figure 29. Crematogaster dalyi [Coonoor, India]. A Lateral view of body B Full face view C Dorsal view
of petiole and postpetiole.

Type material examined. Crematogaster (Oxygyne) dalyi: \ectotype worker (middle
specimen of three on one pin) by present designation and two paralectotype workers
from India, Coonoor (Daly) (MHNG, examined).

Measurements and indices. (type workers, n = 3) HW 0.86-0.92; HL 0.85-0.87;
CI 99-108; SL 0.74-0.75; SI 80-87; EL 0.16-0.18; PW 0.55-0.58; WL 1.02-1.03; PSL
0.1-0.12; PtL 0.25-0.26; PtW 0.30-0.33; PtH 0.16-0.17; PpL 0.17-0.18; PpW 0.37-
0.39; PtHI 64-65; PtWI 115-132; PpWI 217-229; WI 118-126.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two

78 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

teeth smaller. Scape exceeding posterior corner of head, with dense appressed setae,
each of which is less than width of scape. Compound eye slightly projecting beyond
lateral margin of head in full face view.

Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine short and
stout; length longer than spiracle, directed upward; dorsum as high as anterior propo-
deum in lateral view. Propodeal spiracle situated close to propodeal declivity in lateral
view, directed laterally.

In dorsal view, shape of petiole scoop, longer than broad. Petiolar spiracle big, as wide
as half of propodeal spiracle in size. Postpetiolar spiracle big and distinct, located anteriorly
on lateral surface. In dorsal view, postpetiole broader than long, strongly bilobed but with-
out longitudinal median sulcus. Postpetiole distinctly wider than petiole in dorsal view.

Integument essentially smooth and shiny. Clypeus generally smooth and shiny,
but with feable rugulae. Malar region smooth and shiny. Dorsal surface of promesono-
tum smooth and shiny. Lateral surface of pronotum generally smooth and shiny, but
with longitudinal rugulae on anterior portion. Mesopleuron and lateral propodeum
generally smooth and shiny, but with weak longitudinal rugulae. Dorsal surface of
propodeum smooth and shiny.

Erect pilosity absent. Dorsum of head, clypeus and mesosoma with short and ap-
pressed setae. Clypeus with one pair of longer setae on anteriormost portion, directed
medially. Anterior clypeal margin with one single longer setae and two pairs of longer
setae on the median portion, mixed with some shorter setae on sides. No erect seate on
pronotal shoulders. Posterolateral tubercle without appressed setae. Ventral surface of
petiole with appressed setae. Postpetiole without setae posteriorly. Fourth abdominal
tergite with short and appressed setae.

Body color brown.

Distribution. This species is only known from the type locality in India.

Comments. This species is similar to C. sikkimensis, but differs in having short
propodeal spines.

Crematogaster ebenina Forel

Fig. 30

Crematogaster (Oxygyne) ebenina Forel, 1902: 199. Type locality: India, Thana, Kanara,
Poona, Belgaum. Combination in C. (Crematogaster) by Blaimer, 2012b: 55.
Crematogaster ebenina var. corax Forel, 1902: 200. Type locality: Myanmar, Moul-

main. syn. n.

Type material examined. Crematogaster (Oxygyne) ebenina: lectotype worker (bottom
specimen of three on one pin) by present designation and five paralectotype workers
from India, Kanara (Wroughton) (MHNG, examined); one syntype worker from India,
Poona (Wroughton) (NHMW, examined). Crematogaster ebenina vat. corax: six syntype
workers from Myanmar, Moulmain (Hodgson) (MHNG, examined).

Revision of the Crematogaster ranavalonae-group in Asia... 72

Figure 30. Crematogaster ebenina [Gandhi National Park, India]. A Lateral view of body B Full face
view C Dorsal view of petiole and postpetiole.

Non-type material examined. INDIA: 4 workers, Mumbai, 16.x.2007 (S. Hosoishi).

Measurements and indices. (workers, n = 4) HW 0.85-0.94; HL 0.81-0.86; CI
105-109; SL 0.69-0.76; SI 78-82; EL 0.18-0.19; PW 0.51-0.56; WL 0.98-1.05; PSL
0.11-0.13; PtL 0.24-0.26; PtW 0.24-0.27; PtH 0.16-0.17; PpL 0.14-0.16; PpW 0.24-
0.26; PtHI 64-67; PtWI 100-113; PpWI 162-173; WI 96-100.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two

80 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each
of which is less than or as long as width of scape in length. Compound eye large and
slightly projecting beyond lateral margin of head in full face view.

Ventrolateral katepisternal ridge indistinct. Propodeal spine long and stout; length
equal to slightly greater than diameter of propodeal spiracle, directed upward, dorsum
as high as anterior propodeum. Propodeal spiracle large and situated close to propodeal
declivity in lateral view, directed posterolaterally.

In dorsal view, shape of petiole scoop with convex side, longer than broad. Antero-
lateral corner of petiole undeveloped, tapering anteriorly. Petiolar spiracle big, as large as
half of propodeal spiracle in diameter. Postpetiole weakly bilobed but without longitu-
dinal median sulcus. Petiole slightly wider than, or as wide as postpetiole in dorsal view.

Integument essentially smooth and shiny. Clypeus generally smooth and shiny,
but with feable rugulae. Malar region smooth and shiny without rugulae. Dorsal sur-
face of promesonotum smooth and shiny. Lateral surface of pronotum smooth and
shiny. Mesopleuron and lateral propodeum generally shiny, but with feable rugulae.
Dorsal surface of propodeum smooth and shiny.

Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short
and appressed sparse setae. Clypeus without longer setae on anteriormost portion. An-
terior clypeal margin with one single longer setae and two pairs of longer setae, mixed
with some shorter setae on the side. No erect setae on pronotal shoulder. Posterolateral
tubercle with some decumbent to appressed shorter setae. Ventral surface of petiole
with appressed setae. Postpetiole with some shorter setae posteriorly. Fourth abdomi-
nal tergite with appressed setae.

Body color reddish brown.

Distribution. This species is distributed in India and Myanmar.

Comments. This species is unique in Asian C. ranavalonae-group in having a slen-
der petiole. This species is similar to C. tumidula, but distinguished in having the
petiole without anterolateral corners.

The syntype workers of C. ebenina corax agree well with syntype workers of C.
ebenina. Thus I treated C. ebenina corax as a junior synonym of C. ebenina.

Crematogaster hashimi sp. n.
http://zoobank.org/520D 1 15A-F486-43F2-8520-E9114103C9EF
Fig. 31

Type material examined. Holotype. MALAYSIA: Worker from Ulu Gombak, Sel-
angor [N3°19', E101°45'] 15.iii.2010, leg. S. Hosoishi, SH10-Mal-67 (FRIM: KU-
MANTO012). Paratypes. Seven workers, same data as holotype (CASC: KUMAN013;
KUM: KUMANT014; BMNH: KUMANT015, MCZC: KUMANT016; MHNG:
KUMANT017; MBBJ: KUMANT018; THNHM: KUMANT019).

Non-type material examined. THAILAND: Macklong, 29.xi.2003, leg. Watana
Sakchoowong, TUS 15min, WS3 (KUM).

Revision of the Crematogaster ranavalonae-group in Asia... 81

Figure 31. Crematogaster hashimi [Ulu Gombak, Selangor, W. Malaysia]. A Lateral view of body B Full

face view C Dorsal view of petiole and postpetiole.

Measurements and indices. (type and non-type workers, n = 12). HW 0.80-0.88;
HL 0.77-0.82; CI 100-109; SL 0.67-0.75; SI 82-89; EL 0.16-0.18; PW 0.48-0.56;
WL 0.87-0.98; PSL 0.03-0.06; PtL 0.23-0.26; PtW 0.22-0.26; PtH 0.13-0.15; PpL
0.15-0.17; PpW 0.24-0.28; PtHI 54-61; PtWI1 96-109; PpWI 153-167; WI 96-108.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two
teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of
which is less than width of scape in length. Compound eye large and slightly projecting
beyond lateral margin of head in full face view.

82 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine undevel-
oped, but as small tubercule in some workers; length distinctly shorter than propodeal
spiracle. Propodeal spiracle large and circular, situated close to propodeal declivity in
lateral view, directed laterally.

In dorsal view, shape of petiole scoop with convex side, longer than broad. Anterolat-
eral corner of petiole rounded. Petiolar spiracle small, less than half of propodeal spiracle
in size. In dorsal view, postpetiole slightly broader than long. Postpetiole weakly bilobed,
but without longitudinal median sulcus. Postpetiole as wide as petiole in dorsal view.

Integument essentially smooth and shiny. Clypeus smooth and shiny without
rugulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth
and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron smooth and
shiny. Dorsal surface of propodeum smooth and shiny. Lateral propodeum smooth
and shiny.

Erect pilosity almost absent. Dorsum of head, clypeus, mesosoma and fourth ab-
dominal tergite with short and appressed sparse setae . Clypeus without longer setae
on anteriormost portion. Anterior clypeal margin with two or three pairs of longer
setae on median portion, with some shorter setae on side. No erect setae on pronotal
shoulder. Posterolateral tubercle with some appressed setae. Ventral surface of petiole
without setae. Postpetiole without longer erect setae posteriorly. Fourth abdominal
tergite with appressed setae.

Body color reddish brown.

Etymology. The species is dedicated to Dr. Rosli Hashim, University of Malaya,
who helped with field surveys in Malaysia.

Distribution. This species is distributed in Thailand and Malaysia (Peninsula).

Comments. This species is similar to C. imperfecta, but differs in having circular
propodeal spiracles and broader petiole.

This species inhabits well-developed forests, and forage on trees.

Crematogaster imperfecta sp. n.
http://zoobank.org/B9AB38B4-BFOA-44FF-BA07-7E8CFE33FEBB
Fig. 32

Type material examined. Holotype. MALAYSIA: Worker from Sg. Kalang, 800-
1000 m alt., Sabah, Borneo, 23.ii.1997, leg. Sk. Yamane (ITBC: KUMANT020).
Paratypes. Three workers, same data as holotype (CASC: KUMANT021; KUM: KU-
MANT022; SKYC: KUMANT023). Two workers from MALAYSIA: Tower Region,
Lambir NP, Miri, Sarawak, 26.i.1983, leg. Sk. Yamane, Canopy Ecol. (BMNH: KU-
MANT024; THNHM: KUMANT025).

Measurements and indices. (type workers, n = 6). HW 0.77-0.87; HL 0.72-0.81;
CI 103-108; SL 0.63-0.71; SI 78-83; EL 0.16-0.18; PW 0.48-0.54; WL 0.86-0.95;
PSL 0.02-0.04; PtL 0.24-0.26; PtW 0.21-0.23; PtH 0.13-0.15; PpL 0.14-0.16; PpW
0.22-0.26; PtHI 54-60; Pt WI 88-92; PpWI 156-167; WI 104-114

Revision of the Crematogaster ranavalonae-group in Asia... 83

Figure 32. Crematogaster imperfecta [Sg. Kalang, Tenom, Sabah, Borneo, E. Malaysia]. A Lateral view
of body B Full face view C Dorsal view of petiole and postpetiole.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two
teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of
which is less than width of scape in length. Compound eye large and slightly projecting
beyond lateral margin of head in full face view.

Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine undevel-
oped, but as small tubercule in some workers; length distinctly shorter than propodeal
spiracle. Propodeal spiracle large, situated close to the propodeal declivity in lateral
view, directed laterally.

In dorsal view, shape of petiole scoop with convex side, longer than broad. Ante-
rolateral corner of petiole developed. Petiolar spiracle small, less than half of propo-

84 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

deal spiracle in size. In dorsal view, postpetiole slightly broader than long. Postpetiole
weakly bilobed, but without longitudinal median sulcus. Postpetiole as wide as petiole
in dorsal view.

Integument essentially smooth and shiny. Clypeus smooth and shiny without
rugulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth
and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron smooth and
shiny. Dorsal surface of propodeum smooth and shiny. Lateral propodeum smooth
and shiny.

Erect pilosity almost absent. Dorsum of head, clypeus, mesosoma and fourth ab-
dominal tergite with short and appressed sparse setae. Clypeus with longer setae on an-
teriormost portion, directed medially. Anterior clypeal margin with two pairs of longer
setae on median portion, with some shorter setae on side. No erect setae on pronotal
shoulder. Posterolateral tubercle with some appressed setae. Ventral surface of petiole
without setae. Postpetiole without longer erect setae posteriorly. Fourth abdominal
tergite with appressed setae.

Body color reddish brown.

Etymology. The specific name refers to the undeveloped propodeal spines.

Distribution. This species is known from Malaysia (Borneo).

Comments. This species is very similar to C. hashimi, but differs in having ellipti-
cal propodeal spiracles and a slender petiole.

Crematogaster pia Forel
Fig. 33

Crematogaster tumidula subsp. pia Forel, 1911: 384. Type locality: Malaysia, Berhen-
tian Tingi, Nigri Sembilan, Malacca. Combination in C. (Acrocoelia), and raised to
species by Emery, 1922: 151; in C. (Crematogaster) by Blaimer, 2012b: 55.

Type material examined. Crematogaster tumidula subsp. pia: lectotype worker (top
specimen of three on one pin) by present designation and five paralectotype work-
ers from Malaysia, Berhentian Tingi, Nigri Sembilan, Malacca (R. Matrin) (MHNG,
examined); five syntype workers from Malaysia, Berhentian Tingi, Nigri Sembilan,
Malacca (R. Matrin) (NHMB, examined).

Non-type material examined. THAILAND: 4 workers, Maegar, Phayao, 21.iii.1990
(No collector’s name); 6 workers, Thachai Sukhothai, 1.iii.1991 (Vo collector’s name); 21
workers, Ban Krang Camp, Kaeng Krachan National Park, Phet Buri [N12°47', E99°27']
23.xii.2012 (SH12-Tha-01) (S. Hosoishi); 14 workers, Evergreen For., Khlong Naka WS,
Ranong Prov., S. Thailand, 12.viii.2009, (WJT09-TH2045) (W. Jaitrong).

Measurements and indices. (workers, n = 11). HW 0.90-0.96; HL 0.88-0.92; CI
102-108; SL 0.79-0.87; SI 83-94; EL 0.18-0.21; PW 0.54-0.59; WL 1.05-1.13; PSL
0.14-0.17; PtL 0.25-0.27; PtW 0.25-0.30; PtH 0.16-0.18; PpL 0.15-0.19; PpW 0.28-
0.31; PtHI 59-69; PtWI 96-115; PpWI 158-187; WI 103-115.

Revision of the Crematogaster ranavalonae-group in Asia... 85

Figure 33. Crematogaster pia [Ban Krang Camp, Kaeng Krachang National Park, Phet Buri, Thailand].
A Lateral view of body B Full face view C Dorsal view of petiole and postpetiole.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapincal teeth large, basal two
teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of
which is as long as width of scape in length. Compound eye large and slightly project-
ing beyond lateral margin of head in full face view.

Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine long and
stout; length longer than spiracle, dorsum as high as anterior propodeum in lateral

86 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

view; basal width larger than spiracle in diameter. Propodeal spiracle large, situated
close to propodeal declivity in lateral view, directed laterally.

In dorsal view, shape of petiole scoop with convex side, as broad as long. Ante-
rolateral corner of petiole angulate. Petiolar spiracle big, as wide as half of propodeal
spiracle in diameter, directed laterally. In dorsal view, postpetiole broader than long,
bilobed but without longitudinal median sulcus. Postpetiole slightly wider than petiole
in dorsal view.

Integument essentially smooth and shiny. Clypeus smooth and shiny without ru-
gulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth and
shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral propo-
deum generally shiny, but with feable rugulae. Dorsal surface of propodeum smooth
and shiny; anterior dorsum with feable rugulae.

Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short
and appressed sparse setae. Clypeus with one pair of longer setae on anteriormost por-
tion, directed medially. Anterior clypeal margin with two or three pairs of longer setae
on median portion, mixed with some shorter setae on side. No erect setae on prono-
tal shoulder. Posterolateral tubercle with some decumbent to appressed shorter setae.
Ventral surface of petiole with appressed setae. Postpetiole with some shorter setae
posteriorly. Fourth abdominal tergite with appressed setae.

Body color reddish brown to brownish.

Distribution. This species is distributed in Thailand and Malaysia (Peninsula).

Comments. This species is similar to C. butteli and C. tumidula, but differs from
C. butteli in having an anterolaterally developed petiole, and from C. tumidula in hav-
ing long propodeal spines.

This species inhabits well-developed forests, make a carton nest and forage on trees.

Crematogaster sikkimensis Forel, stat. n.

Fig. 34

Crematogaster (Oxygyne) dalyi var. sikkimensis Forel, 1904: 24. Type locality: India,
Sikkim.

Type material examined. Crematogaster (Oxygyne) dalyi var. sikkimensis: lectotype
worker (top specimen of three on one pin) by present designation and three paralecto-
type workers from India, Sikkim (W6/ler) (MHNG, examined).

Measurement and indices. (type workers, n = 4). HW 0.82-0.91; HL 0.80-0.88;
CI 102-103; SL 0.72-0.78; SI 83-89; EL 0.17-0.19; PW 0.50-0.56; WL 0.92-1.07;
PSL 0.14-0.17; PtL 0.25-0.26; PtW 0.27-0.30; PtH 0.14-0.17; PpL 0.14-0.17; PpW
0.33-0.36; PtHI 60-65; PtWI 108-116; PpWI 212-236; WI 120-130.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapincal teeth large, basal two
teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of

Revision of the Crematogaster ranavalonae-group in Asia... 87

Figure 34. Crematogaster sikkimensis [Sikkim, India]. A Lateral view of body B Full face view C Dorsal
view of petiole and postpetiole.

which is as long as width of scape in length. Compound eye large and slightly project-
ing beyond lateral margin of head in full face view.

Ventrolateral katepisternal ridge appearing indistinct posteriorly. Propodeal spine
long and slender; length longer than spiracle, directed upward; dorsum higher than
anterior propodeum. Propodeal spiracle large, situated close to propodeal declivity in
lateral view, directed laterally.

In dorsal view, shape of petiole scoop with convex side, as long as broad. Petiolar
spiracle big, as wide as half of propodeal spiracle in diameter. In dorsal view, postpe-
tiole broader than long, strongly bilobed laterally, but without longitudinal median
sulcus. Postpetiole wider than petiole in dorsal view.

88 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

Integument essentially smooth and shiny. Clypeus smooth and shiny without ru-
gulae. Malar region with feable rugulae. Dorsal surface of promesonotum smooth and
shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral pro-
podeum generally shiny, but with longitudinal rugulae. Dorsal surface of propodeum
smooth and shiny.

Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short
and appressed sparse setae. Clypeus with one pair of longer setae on anteriormost por-
tion, directed medially. Anterior clypeal margin with two pairs of longer setae, mixed
with some shorter setae on side. No erect setae on pronotal shoulder. Posterolateral
tubercle with some decumbent to appressed shorter setae. Ventral surface of petiole
with appressed setae. Postpetiole with some shorter setae posteriorly. Fourth abdomi-
nal tergite with appressed setae.

Body color brown.

Distribution. This species is only known from the type locality in India.

Comments. This species is similar to C. daisyi and C. dalyi, but differs from C.
daisyi in having the spiracles on the petiole located on the middle position in lateral
view, and from C. dalyi in having long propodeal spines.

Top specimen of three on one pin was designated as lectotype worker here, but it
is noted that the body of the middle specimen was lost from the card point.

Crematogaster tumidula Emery
Fig. 35

Crematogaster tumidula Emery, 1900: 689, fig. 8. Type locality: Indonesia, Pangherang-
Pisang, Sumatra. Combination in C. (Physocrema) by Emery, 1922: 140; Don-
isthorpe 1941: 226; in C. (Oxygyne) by Hosoishi & Ogata, 2009: 33; in C. (Cre-
matogaster) by Blaimer, 2012b: 55.

Type material examined. Crematogaster tumidula: \ectotype worker by present desig-
nation and one paralectotype worker from Indonesia, Pangherang-Pisang, Sumatra (E.
Modigliani) (MCSN) [examined].

Non-type material examined. MALAYSIA: 2 workers, Old Tower R., Lambir N.
P., Miri, Sarawak, 30.xii.1997, (SR97-SKY-106) (S&. Yamane); 3 workers, Bt. Pantu,
Lambir N. P., Miri, Sarawak, 13.viii.1995, (Sk. Yamane); 1 worker, Tower Region,
Lambir N. P., Miri, Sarawak, 21.1.1993, (Sk. Yamane); INDONESIA: 1 worker,
Sangkimah, Kutai N. P., Kalimantan, 17.ix.1993, (Sk. Yamane).

Measurements and indices. (workers, n = 10). HW 0.73-0.88; HL 0.72-0.86; CI
101-107; SL 0.60-0.77; SI 80-88; EL 0.15-0.19; PW 0.43-0.53; WL 0.80-0.97; PSL
0.08-0.12; PtL 0.20-0.26; PtW 0.23-0.27; PtH 0.14-0.18; PpL 0.14-0.22; PpW 0.20-
0.28; PtHI 54-68; PtWI 89-113; PpWI 125-178; WI 88-118.

General description of worker. Head appearing subquadratic in front view.
Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two

Revision of the Crematogaster ranavalonae-group in Asia... 89

Figure 35. Crematogaster tumidula [Lambir National Park, Sarawak, Borneo, E. Malaysia]. A Lateral
view of body B Full face view C Dorsal view of petiole and postpetiole.

teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of
which is less than width of scape in length. Compound eye large and slightly projecting
beyond lateral margin of head in full face view.

Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine short and
stout; length as long as spiracle, directed posteriorly, dorsum as high as anterior propo-
deum; basal width smaller than spiracles in diameter. Propodeal spiracle large, situated
close to propodeal declivity in lateral view, directed laterally or posterolaterally.

90 Shingo Hosoishi / Journal of Hymenoptera Research 42: 63-92 (2015)

In dorsal view, shape of petiole scoop with convex side, longer than broad. An-
terolateral corner of petiole developed. Petiolar spiracle as large as half of propodeal
spiracle in diameter, directed laterally. Postpetiole bilobed but without longitudinal
median sulcus. Postpetiole as wide as petiole in dorsal view.

Integument essentially smooth and shiny. Clypeus smooth and shiny without ru-
gulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth and
shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral propo-
deum generally smooth, but with feable rugulae. Dorsal surface of propodeum smooth
and shiny; anterior dorsum smooth without rugulae.

Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short and
appressed sparse setae. Clypeus with one pair of longer setae on anteriormost portion, di-
rected medially. Anterior clypeal margin with two pairs of longer setae on median portion,
mixed with some shorter setae on side. No erect setae on pronotal shoulder. Posterolateral
tubercle with some appressed shorter setae. Ventral surface of petiole with appressed setae.
Postpetiole with some appressed setae. Fourth abdominal tergite with appressed setae.

Body color reddish brown.

Distribution. This species is known from Malaysia (Borneo) and Indonesia (Ka-
limantan, Sumatra).

Comments. This species is similar to C. ebenina, but differs in having developed
anterolateral corners of the petiole. It is also similar to C. pia, but differs in having
short and stout propodeal spines and the basal portion with smooth surface.

Crematogaster vandermeermohri Menozzi

Crematogaster (Paracrema) vandermeermohri Menozzi, 1930: 1, fig. 1. Type locality:
Indonesia, Toetoepan, Sumatra. Combination in C. (Crematogaster) by Blaimer,

2012b: 55.

Type material. Crematogaster (Paracrema) vandermeermohri: syntype worker from In-
donesia, Toetoepan, Sumatra (probably in IEGG, not seen).

Comments. | have not been able to examine type-material of C. vandermeermohri.
Judging from the original description and figure, this species is close to C. hashimi and
C. imperfecta. Menozzi (1930) described the following features: (1) propodeal spines
small and blunt, inflated near the base; (2) postpetiole slightly narrower than petiole in
dorsal view. The two features are not found in the related species. The postpetiole is as
wide as or wider than petiole in C. hashimi and C. imperfecta.

Acknowledgments

I would like to thank Kazuo Ogata and Seiki Yamane for collecting specimens and
their continuing support. I also thank to the following persons for loans of material

Revision of the Crematogaster ranavalonae-group in Asia... 91

or access to museum collections: Daniel Burckhardt (NHMB); Bernhard Merz
(MHNG); Maria Tavano and Roberto Poggi (MCSN); Suzanne Ryder and Gavin
Broad (BMNH); Dominique Zimmermann (NHMW). Barry Bolton provided a
suggestion on the terminology of katepisternum. Additional material was received
from Watana Sakchoowong, Weeyawat Jaitrong, Katsuyuki Eguchi. Thanks are also
for due to Rosli Hashim and Wattanachai Tasen who provisioned my collecting trips.
I would like to thank the ANeT members for encouragement. Thanks are also due
to Mark Lorenz (Forte Inc.) for improving the English. This work was supported in
part by JSPS KAKENHI (Grant-in-Aid for Scientific Research (C)) Grant Number
26440221.

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