D. W. Davidson, S. C. Cook, R. R. Snelling and T. H.

Chua
“Explaining the Abundance of Ants in Lowland Tropical Rainforest Canopies”

Supporting Online Material

1. Materials and Methods

Arthropods were collected haphazardly from foliage or leaf litter and stored in silica gel or 70%

ETOH for transport to Utah. Leaf tissue (~ 10 cm2 and collected from haphazardly chosen
plants) was air-dried in a parabolic solar oven (Peru) or dry room (Brunei) and stored in

desiccant. Except for the smallest arthropod samples (~ 0.600 mg, analyzed whole), dry samples

were ground to powder with a modified pipette tip in liquid N in 1.5 ml microcentrifuge tubes.

On a Sartorius MC-5 microbalance, 0.600 to 1.200 mg dry samples (or ~ 2.000 mg for plants)

were weighed into 3.5 x 5 mm tin cups (Costech Analytical Technologies, Inc. Valencia CA,

#041060) and stored in a desiccator until analyzed in the University of Utah’s SIRFER facility

on a Finnigan Delta S isotope ratio mass spectrometer (Bremen, Germany) or a Finnigan Delta

Plus IRMS.

In nitrogen (N) isotope analyses, ∗15N (‰) = [(Rsample / Rstandard) -1 x 1000], where R =
15
N/14N ratio, and the standard is calibrated to NIST international standards. We assume that ants

feeding at different trophic levels allocate dietary N in similar ways across similar distributions

of tissue types. Carbon isotope ratios, produced in these same analyses, increased as expected

(S1) with ∗15N ‰, but there was much scatter due to variation in canopy height (S2) and other

factors, and we present these data elsewhere.

Figs. 1 & 2 report predominant activities by species and category (authors’ unpublished

data): trophobiont-tending, leaf-foraging, and neither or poorly known (authors’ unpublished

data). Independence of data was promoted by either censussing ant activities just once along

particular trails in a given year (Peru, 2000-2001), or repeating censuses along the same trail

during different days, but purposefully avoiding replicate observations on the same plant

(Brunei, 2001-2002, with a more limited trail system). The latter method was unlikely to have

compromised the integrity of our results, because prodigious tending of trophobionts appeared to
be a species-level trait, and there was little intraspecific variation among colonies. Data are

supplemented for some species from earlier field notes (D. Davidson, unpubl. data, Peru) and

literature reports of ant activities in the high canopy (Table S1). Ants foraging in leaf litter were

assumed to be hunting or scavenging prey.

2. Supporting Text

2.1 Study sites, collections and vouchers

Study sites were: Estación Biológica de Cocha Cashu, Parque Nacional Manu, Madre de Dios,

Peru, 11o54'S 71o22'W, ~350 m, mean annual precipitation = ~ 2.2 m (highly seasonal), lowland

moist forest (S3), and Kuala Belalong Field Studies Centre, Batu Apoi Forest Reserve,

Temborong District, Brunei Darussalam, 4o 32', 115° 10'E, 200 m, mean annual precipitation >

4,000 mm (largely aseasonal), lowland mixed dipterocarp forest (S4). Collections were made in

September through mid-November, 1998-2001 (transition to wet season), in Peru, and May -

July, 2001-2002 (onset of brief dry periods), in Brunei. Voucher specimens of ants have been

deposited in collections of the Entomology Division, Natural History Museum of Los Angeles

County (most species) and either Peru’s Museo de Historia Natural (1998 - 99) and the Museo de

Entomología, Universidad Nacional Agraria La Molina, Lima, Peru (2000 - 01), or the Brunei

Natural History Museums, Bandar Seri Begawan, Brunei. Unicate samples, including those of
ants, all trophobionts and arthropod herbivores and predators, were destroyed in isotopic

analyses after identification. Plants (Table S1) were identified by P. Nuñez and vouchered at the

herbarium of the Universidad San Antonio Abad en Cusco, or compared with published accounts

and drawings (Brunei).

2.2 Site differences in isotopic ratios

We cannot say for certain why isotopic ratios were lower on average in Brunei than in Peru, but

we suspect that latitudinally-determined differences in amount and seasonality of precipitation

(above), together with disparities in topography and soil fertility, could have affected the

2
availability of different N sources to both producers and consumers. Our data are consistent with

the balance of N from precipitation, versus animal excretion and decay of plant and animal

biomass, being greater in Brunei than in Peru. Some species or genera sampled at the beginning

of the wet season in Peru exhibited substantially lower values when collected at the end of the

wet season in Panama (S5).

2.3 Variation in isotopic ratios within ant subfamilies

Ponerinae and Ecitoninae. Among ponerines, values for two well-known exudate-feeders (nos.
2,6, fig. 1) were comparatively low but still identified these species as predators/scavengers of N.

Within army ants, ∗15N was lower for specialized predators of low ratio formicine and

dolichoderine ants or leafcutter ants (ecitonines 1 and 2, respectively), than for species using

mass foraging to flush and capture (mainly predatory) leaf litter arthropods [ecitonines 3 and 4,

(S6), Table S1].

Dolichoderinae. Exceptionally carnivorous Peruvian dolichoderines included three specialized

Azteca plant-ants (fig. 1, 16 - 18) and some free-living, trophobiont-tending Azteca (14,15,19-

21). Behavioral assays are available for the majority of these species and invariably show severe

protein-limitation (authors’ unpublished data). Associated with immobile Coccidae on stems and
trunks, these species may obtain poorer quality honeydew (S7) than do ants herding mobile

membracids to high-N feeding sites on young leaves and reproductive pedicels (S8, S9).

Exhibiting somewhat lower values in relation to other Azteca plant-ants were two species (nos. 6,

8) from legume hosts, which can themselves average relatively low ∗15N (S10) values.

Myrmicinae. Exceptionally predacious Crematogaster species included the only Peruvian leaf-

litter forager (myrmicine 24), and two Bruneian species (nos. 18, 27).

The highest individual ratio for Bruneian Crematogaster, and among the highest values for

Peruvian myrmicines (Wasmannia, species 31) are “unicolonial” taxa (superscript “U”) forming

3
“supercolonies” in their native or introduced ranges, respectively (S11, see also Linepithema

humile - dolichoderine 22, fig. 1). These species forage extensively for exudates, but obtain N

mainly from prey; abundant dietary carbohydrates (CHOs) may subsidize large colonies of tiny,

pugnacious workers (S12, S13).

2.4 Herbivory and its correlates

Although isotopic evidence is consistent with herbivory contributing substantial N to

many arboreal exudate-foragers, a contrarian might object that organisms feeding principally as
mutualists of either EFN-producing plants or herbivorous trophobionts are not technically

herbivores. However, significant fractions of N [and probably carbon] pools of exudate-feeding

ants are acquired without breakdown of herbivore tissues, and exudate-feeding ants have

unbalanced diets typical of herbivores (S14, authors’ unpublished data). CHOs in excess of

amounts paired with N for growth and reproduction may enhance N capture when deployed to

support N-contributing symbionts (S15) or large (competitive) colonies of mainly CHO-

dependent workers (S16), or to fuel rapid foraging tempos, territoriality, and offensive and

defensive weaponry (S14). By funding energetically expensive search for prey, they may also

have facilitated a move to the arboreal zone, where prey densities are likely lower than on the

two-dimensional forest floor.

2.5 Effects of ants on plants

Taxa obtaining substantial N from prey are most likely to benefit plants by reducing herbivore

loads. However, residence times of legionary taxa are too brief to provide significant protection,

and except for Paraponera clavata and a few Pachycondyla species, most non-legionary

predators [figs. 1 & 2, Table S1] forage terrestrially or in low vegetation. In contrast, as a group,

leaf-foragers occupy the full canopy volume, frequent EFNs, search foliage continuously and

intensively when active, and take prey opportunistically (S17, authors’ unpublished data). These

species may regularly benefit a diversity of plants, though indirect effects of these ants on plants

4
(e.g., those mediated via interactions with epiphylls) are currently poorly understood.

Our findings suggest that high ∗15N values may predict which ant species are likely to be
useful in suppressing herbivores in agricultural settings. However, it is important not to overlook

highly carnivorous taxa with intermediate isotopic signatures due to the consumption of low ratio

prey. Dolichoderus thoracicus (fig. 2, dolichoderine 4), the most protein-starved species in our

Bruneian sample (authors’ unpublished data), may represent such a taxon (e.g., S18).

3. Supplementary Table S1
Tables S1A (Peru) and S1B (Brunei) present ant identifications, samples sizes, and natural

history notes. Nomenclature follows Bolton (S19), but Dendromyrmex is recognized as a sub-

genus of Camponotus (S20), all sampled cephalotines are classified as Cephalotes (S21), and

several new species of Pheidole are included (S22). Unidentified Camponotus species are

tentatively assigned to subgenera; identifications (including subgenera) of Polyrhachis are

courtesy of R. J. Kohout.

Table S1A Identifications (by subfamily) and sample sizes (N) of ants in fig. 1
(Peru)

Subfamily ID Species (sub-genera in N Host plant of plant-

no. parentheses) ants, or other natural
history notes
Formicinae 1 Camponotus novogranadensis 1
2 Camponotus iheringi 1
3 Camponotus heathi 1
4 Camponotus (Myrmaphaenus) sp. B 6
5 Camponotus (Dendromyrmex) sp. A 2
6 Camponotus (Myrmobrachys) sp. A 7
7 Camponotus rectangularis 1
8 Camponotus sexguttatus 2
9 Camponotus (Myrmaphaenus) sp. C 3
10 Camponotus (Colobopsis) sp. 2
11 Camponotus (Myrmobrachys) sp. B 3

5
 12 Camponotus (Myrmobrachys) sp. C 1
13 Camponotus (Dendromyrmex) sp. B 2
14 Camponotus (Myrmomalis) 4 Guadua spp. (dead
depressus bamboo culms), feed
on insect secretions at
culm tips*
15 Camponotus longipilis 4 Guadua spp., tend
Coccidae in live
bamboo culms*
16 Camponotus (Dendromyrmex) 3
nidulans
17 Camponotus branneri 1
18 Camponotus sericeiventris 7 inhabit live
trunks/branches of a
variety of tree species
19 Gigantiops destructor 7
20 Camponotus femoratus 9 ant-garden epiphytes
(S9)
21 Camponotus mirabilis 3 Guadua spp., tend
Coccidae in live
bamboo culms; mass
forage for terrestrial
prey*
22 Camponotus atriceps 1 highly predatory
nocturnal species; tend
membracids (S23)
23 Paratrechina sp. 3
24 Myrmelachista sp. 5 Pleurothyrium sp., ants
tending (and
consuming?)
Coccidae within stems

Dolichoderinae 1 Azteca ovaticeps 9 Cecropia

membranacea (S24)
2 Dolichoderus cf. bispinosus A 8
3 Dolichoderus quadridenticulatus 6
4 Dolichoderus lugens 5
5 Dolichoderus cf. bispinosus B 6
6 Azteca sp. A 1 unidentified legume
host
7 Dolichoderus debilis 9

6
 8 Azteca sp. B 4 Lonchocarpus sp.
(legume)
9 Dolichoderus bidens 22
10 Dolichoderus attelaboides 9
11 Dolichoderus decollatus 8
12 Azteca sp. C 3 free-living, carton-
builder
13 Azteca sp. D 1 free-living, nest?
14 Azteca sp. E 2 free-living, cavity-
nester
15 Azteca sp. F 4 free-living, carton-
builder
16 Azteca sp. G 5 from Triplaris
poeppigiana (S25)
17 Azteca sp. H 7 Nectandra sp.
18 Azteca sp. I 7 Cordia nodosa, carton-
builder (S25)
19 Azteca sp. J 2 ant-garden epiphytes
(S9)
20 Azteca sp. K 1 free-living, carton-
builder, associate of
Dolichoderus
decollatus
21 Azteca sp. L 1 free-living cavity-
nester, associate of
Do. decollatus
22 Linepithema humile 1 unicolonial (forming
'supercolonies') in
introduced range (see
main article)

Myrmicinae 1 Cephalotes sp. A 1 limited activity data

2 Cephalotes cordatus 2 limited activity data
3 Cephalotes sp. B 1 limited activity data
4 Cephalotes clypeatus 1 limited activity data
5 Cephalotes oculatus 3
6 Cephalotes umbraculatus 1
7 Cephalotes atratus 3 high specificity in
trophobiont relations
(S23)
8 Crematogaster stollii 1
9 Daceton armigerum 1

7
10 Atta columbica 4 cultivate fungi on fresh
vegetation (S11)
11 Crematogaster carinata 12
12 Crematogaster brasiliensis 7
13 Crematogaster arcuata 2
14 Crematogaster limata 5
15 Pheidole minutula 2 Clidemia heterophylla
and Maieta guianensis
(S25)
16 Crematogaster laevis 1 Clidemia heterophylla
and Maieta guianensis
(S25)
17 Crematogaster acuta 6
18 Crematogaster levior 4 ant-garden epiphytes
19 Ochetomyrmex sp. 9
20 Allomerus sp. 5 Cordia nodosa (S25)
21 Cyphomyrmex cf. major 1 cultivate fungi on
feces, corpses, dead
fruit (S11)
22 Pheidole longiseta 1 new sp. (S22)
23 Acromyrmex sp. 1 cultivate fungi on fresh
vegetation (S11)
24 Crematogaster nigropilosa 1 nesting terrestrially,
foraging in leaf litter
25 Pheidole biconstricta 6
26 Crematogaster egregior 1
27 Pheidole carapuna 2
28 Pheidole fissiceps 3 new sp. (S22)
29 Wasmannia sp. A 1 EFN and trophobionts
but few observations
30 Pheidole sp. nov. 1 1 new sp. (S22)
31 Wasmannia auropunctata 5
32 Solenopsis parabiotica 1 brood parasite of ant-
garden ants (S9)
33 Apterostigma cf. pilosum 1 cultivate fungi on
insect feces, dead
wood (S11)
34 Pheidole minax 1 new sp. (S22)
35 Wasmannia sp. B 1
36 Apterostigma sp. A 1 see A. cf. pilosum

8
Ponerinae 1 Pachycondyla luteola 2 Cecropia sp. nov.
(S25)
2 Paraponera clavata 6 nest terrestrially,
forage in high canopy
3 Gnamptogenys annulata 3 nest in low arboreal
zone
4 Pachycondyla cf. oberthueri 2 nest in low arboreal
5 Pachycondyla sp. A 1 nest arboreally
6 Ectatomma tuberculatum 10 nest terrestrially,
forage mainly in low
vegetation
7 Platythyrea angusta 2 nest in low arboreal
zone; common in live
Guadua spp. Bamboo*
8 Pachycondyla unidentata 8 cavity nester in low
arboreal zone
9 Pachycondyla villosa 4 nests terrestrially and
arboreally in dead
wood; forages
arboreally
10 Odontomachus sp. 1
11 Anochetus sp. 1
12 Pachycondyla sp. B 1
13 Gnamptogenys sp. (striatula group) 2
14 Pachycondyla apicalis 8 nests terrestrially,
forages terrestrially
and in low arboreal
15 Pachycondyla crassinoda 1
16 Odontomachus haematodus 1 nests terrestrially in
dead wood
17 Gnamptogenys pleurodon 1
18 Gnamptogenys concinna 1 EFN forager and
ambush predator of
bees (S26)
19 Gnamptogenys triangularis 1 millipede predator
(S27)
20 Pachycondyla commutata 6 predator of termites
eating leaf litter (S28)
21 Gnamptogenys horni 1 predator of leaf-litter
ants and other
arthropods (S29)

9
Ecitoninae 1 Eciton hamatum 6 brood predator of
(esp.) Dolichoderus,
formicines, Atta, wasps
(16)
2 Nomamyrmex esenbeckii 2 specialized brood
predator of Atta and
other ants (S30)
3 Eciton burchellii 6 generalized predator of
arthropods, including
immature stages of
social insects (S6)
4 Labidus praedator 4 diet similar to that of E.
burchelli (S6)

Pseudomyrmecinae 1 Pseudomyrmex gracilis 1

2 Pseudomyrmex filiformis 1
3 Pseudomyrmex tenuis 1
4 Pseudomyrmex dendroicus 8 Triplaris americana
(S25n)
5 Pseudomyrmex sp. 1

* D. W. Davidson, J. A. Arias, J. Mann, unpublished data

Table S1B Identifications (by subfamily) of ants in fig. 2 (Brunei):

Subfamily ID Species (subgenera in N Host plant of plant-

no parentheses) ants, or other natural
Formicinae 1 Cladomyrma dianae 3 Neonauclea sp. (S31)
2 Cladomyrma cryptata 1 Milletia nieuwenhuisii
(legume), tends
Pseudococcidae (S31)
3 Camponotus cylindricus 2
4 Echinopla sp. nov. 1
5 Camponotus sp. C 1
6 Polyrhachis (Myrma) rixosa 2
7 Polyrhachis (Myrmhopla) cephalotes 1
8 Camponotus saundersi (cylindricus 4 trophobionts in high
group) canopy cannot be
ruled out.
9 Polyrhachis (Myrmhopla) nr. pressa 2

10
 10 Camponotus sp. BA (cylindricus 3 trophobionts in high
group) canopy cannot be
ruled out
11 Camponotus sp. Y (cylindricus group) 3
12 Camponotus sp. DR (cylindricus 2
group)
13 Camponotus sp. D 1
14 Polyrhachis (Polyrhachis) olybria 1
15 Echinopla melanarctos 6
16 Polyrhachis (Myrma) beccarii 1
17 Echinopla pallipes 2
18 Polyrhachis (Myrma) illaudata 6
19 Polyrhachis (Myrma) nigropilosa 3
20 Camponotus clerodendri (cylindricus) 3 high canopy
group) trophobiont-tending not
ruled out
21 Camponotus sp. B 1
22 Polyrhachis (Myrma) sculpturata 6
23 Polyrhachis (Myrmhopla) armata 5 limited activity data
24 Camponotus festinus 2
25 Camponotus horrens 2
26 Polyrhachis (Polyrhachis) ypsilon 1
27 Polyrhachis (Myrma) sp. nov. 5 (R. J. Kohout, mss)
28 Camponotus sp. BLK (cylindricus 1
group)
29 Camponotus camelinus 5
30 Polyrhachis (Polyrhachis) bihamata 9 limited activity data
31 Camponotus sp. E 1
32 Camponotus gigas 3 predatory trophobiont-
tender (S32)
33 Camponotus sp. F 1 Calamus sp.
34 Camponotus sp. A 1
35 Paratrechina metallescens 3
36 Oecophylla smaragdina 6

Dolichoderinae 1 Dolichoderus sp. 1 2

2 Dolichoderus cuspidatus s. l. 5
3 Dolichoderus cf. affinis 3
4 Dolichoderus thoracicus 10

11
 5 Technomyrmex albipes 2

Myrmicinae 1 Crematogaster sp. A (decamera 1 Macaranga hosei

group) (S33)
2 Cataulacus sp.3 2
3 Crematogaster cf. borneensis A 1 Macaranga winkleri
(S33)
4 Cataulacus horridus 10
5 Crematogaster sp. 5 1
6 Crematogaster cf. borneenis B 1 Macaranga sp.
7 Crematogaster sp. 4 2
8 Crematogaster cf. decamera 4 Macaranga beccariana
(S33)
9 Crematogaster difformis 1 high canopy
trophobiont-tending not
ruled out
10 Crematogaster cf. ebenina 1
11 Crematogaster sp. 2 1
12 Myrmicaria melanogaster 2
13 Meranoplus castaneus 1
14 Crematogaster sp. 3 2
15 Cataulacus latissimus 1
16 Pristomyrmex bicolor 1
17 Myrmicaria brunnea 4
18 Crematogaster sp. 1 1
19 Vollenhovia sp. brevicornis group 1
20 Lophomyrmex bedoti 1
21 Pheidole longipes 1
22 Myrmicaria cf. lutea 1
23 Myrmicaria sp. brunnea group 2 limited activity data
24 Vollenhovia rufiventris 1
25 Pheidole clypeocornis 1
26 Pheidole cariniceps 1
27 Crematogaster sp. 6 1 unicolonial, authors'
observations
28 Pheidologeton sp. 1
29 Pheidole sp. C 1
30 Pheidole plagiaria 2

12
Ponerinae and 1 Myopias modiglianii 1
Aenictinae 2 Diacamma rugosum 4 arboreal forager
3 Pachycondyla cf. tridentata 1
4 Leptogenys kitteli 1
5 Gnamptogenys menadensis 1 leaf-forager
6 Diacamma intricatum 5 terrestrial forager
7 Aenictus gracilis 2
8 Odontoponera transversa 1
9 Leptogenys birmana 3
10 Anochetus sp. nov. 2
11 Leptogenys crassicornis 1
12 Leptogenys chalybaea
Pseudomyrmecinae 1 Tetraponera attenuata 5

4. References and Notes

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the Tropics, Vol. 2, P. Jaisson, Ed. (Université Paris-Nord, Paris, 1983), pp. 59-73.

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8. R. C. Buckley, Ann. Rev. Ecol. Syst. 18, 111 (1987).

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13
11. B. Hölldobler, E. O. Wilson, The Ants (Belknap Press, Harvard Univ., Cambridge MA,

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(Harvard University Press, Cambridge, MA, 2003), 818 pp.

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14
30. M. B. Swartz, Biotropica 30, 682 (1998).

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33. Various undescribed but host-specific species of “Crematogaster decamera” and

“Crematogaster borneensis” [T. Itino et al., Ecol. Res. 16, 787 (2001)] are kept separate

here.

15