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Nest materials and some chemical
characteristics of nests of a New World
swarm-founding polistine wasp, Polybia
paulista (Hymenoptera Vespidae)
K. Kudô a , Sô. Yamane b , S. Mateus c , K. Tsuchida a , Y. Itô d , S. Miyano
e , H. Yamamoto f & R. Zucchi c
a Laboratory of Entomology, Faculty of Agriculture, Gifu University, Gifu,
501-1193, Japan E-mail:
b Biological Laboratory, Faculty of Education, Ibaraki University, Mito,
310-8512, Japan
c Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de
Ribeirão Preto, Universidade de São Paulo, USP, São Paulo, 14040-901,
Brazil
d 2908, Kaguyama 2-chome, Nisshin, Aichi, 470-0134, Japan
e Natural History Museum and Institute, 955-2 Aoba-cho, Chuo-ku, Chiba,
260-8682, Japan
f Chemical Laboratory, Faculty of Education, Ibaraki University, Mito,
310-8512, Japan
Published online: 19 May 2010.
To cite this article: K. Kud , S. Yamane , S. Mateus , K. Tsuchida , Y. It , S. Miyano , H. Yamamoto & R.
Zucchi (2001) Nest materials and some chemical characteristics of nests of a New World swarm-founding
polistine wasp, Polybia paulista (Hymenoptera Vespidae), Ethology Ecology & Evolution, 13:4, 351-360
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Ethology Ecology & Evolution 13: 351-360, 2001
Nest materials and some chemical characteristics of
nests of a New World swarm-founding polistine wasp,
Polybia paulista (Hymenoptera Vespidae)
2
3
1
4
5
K. KUDÔ 1,7, SÔ. YAMANE , S. MATEUS , K. TSUCHIDA , Y. ITÔ , S. MIYANO ,
6
3
H. YAMAMOTO and R. ZUCCHI
1
Laboratory of Entomology, Faculty of Agriculture, Gifu University, Gifu 501-1193,
Japan (E-mail: kudok@cc.gifu-u.ac.jp)
2
Biological Laboratory, Faculty of Education, Ibaraki University, Mito 310-8512,
Japan
3
Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão
Preto, Universidade de São Paulo, USP, São Paulo 14040-901, Brazil
4
2908, Kaguyama 2-chome, Nisshin, Aichi 470-0134, Japan
Natural History Museum and Institute, Chiba, 955-2 Aoba-cho, Chuo-ku, Chiba
5
260-8682, Japan
6 Chemical Laboratory, Faculty of Education, Ibaraki University, Mito 310-8512, Japan
Received 5 January 2001, accepted 5 June 2001
Nest material, nitrogen and amino acid composition of nests were exam-
ined in a New World swarm-founding polistine wasp, Polybia paulista. This wasp
used minute vegetable chips, which were a dominant material, plant hairs, and
mud and/or inorganic particles as nest building materials. A SEM observation
and nitrogen and amino acid contents indicated that the amount of oral secre-
tion used for construction and maintenance of the nests was quite small, com-
pared with that used in Polistes nests. Such a small amount of oral secretion
used for nest building is considered to be associated with their social organiza-
tion (division of labor) and nest material. This study hypothesized that the
amounts of oral secretion used for nest building are determined by an interac-
tion among social organization, nest material and environmental factors, such as
precipitation.
Twenty-four amino acids were detected from protein in the nests of P.
paulista, of which serine, glycine, alanine, valine, proline, aspartic acid and glu-
tamic acid were major components. Amino acid composition of the protein in
the nests of P. paulista differed distinctly from those of other so far known polis-
tine and vespine wasps. The present result supports the view that amino acid
composition of the protein in nests reflects phylogenetical relationships among
wasps.
KEY WORDS: social wasp, Polybia paulista, swarm-founding, nest material, oral
secretion, amino acid composition.
7 To whom correspondence should be addressed.

352
K. Kudô et alii
Introduction
Materials and methods
Results
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Fine surface structure
Nest materials
Oral secretion
Nitrogen content
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Amino acid composition of protein in the envelope and cell walls
Discussion
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Nest materials
Factors determining the amount of oral secretion used for nest building
Amino acid composition of protein in envelope and cell walls
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Acknowledgments
References
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INTRODUCTION
Social wasp nests serve as a place for rearing brood and the centre for their
nesting activities (JEANNE 1977, STARR 1991). Wasps collect woody fibres, minute
vegetable chips, plant hairs and mud, which they masticate and mix with oral
secretion to construct the nest with a variety of architectural design (JEANNE 1975,
WENZEL 1991). This salivary secretion is also used to physically maintain their
nests. Wasps frequently smear the surface of their nests with the secretion to pro-
tect the nest from rain and weathering. It has been reported recently in two Japan-
ese Polistes wasps, P. chinensis and P. riparius that a large amount of the secretion
is used for construction and maintenance of pre-emergence nests (KUDÔ et al. 1998,
YAMANE et al. 1998, KUDÔ 2000).
Recently, chemical characteristics of oral secretions have been examined in
some genera of the subfamily Polistinae, Protopolybia, Ropalidia and Polistes
(SCHREMMER et al. 1985; MCGOVERN et al. 1988; MASCHWITZ et al. 1990; ESPELIE &
HIMMELSBACH 1990; SINGER et al. 1992; KUDÔ et al. 2000a, 2000b; see a review by
JEANNE 1996). Most of these studies indicate that the secretion is primarily com-
posed of protein. MASCHWITZ et al. (1990) and KUDÔ et al. (1998) estimated in Ropa-
lidia opifex and Polistes chinensis that the protein content contained in the oral
secretion was about 90% and 70%, respectively. On the contrary, SCHREMMER et al.
(1985) detected chitin in the nest of swarm-founding polistine wasp of the New
World, Protopolybia chartergoides (cited as Pseudochartergus chartergoides). Since
the oral secretion is composed primarily of protein in most wasp genera so far ana-
lyzed (JEANNE 1996), it is presumed that the cost for secretion production is very
high (MASCHWITZ et al. 1990, KUDÔ et al. 1998, KUDÔ 2000).
The amino acid composition of protein in the nests has been examined in
Dolichovespula maculata (MCGOVERN et al. 1988), R. opifex (MASCHWITZ et al. 1990),
Polistes annularis (ESPELIE & HIMMELSBACH 1990), P. metricus (SINGER et al. 1992),
P. riparius (KUDÔ et al. 2000a) and P. chinensis (KUDÔ et al. 2000b). These results
showed that alanine, glycine and serine were major components in the nests of
these species. Comparing the amino acid composition among the above species, it
was found that the composition was closely similar among Polistes species, but dis-
tinctly differed among different genera (KUDÔ et al. 2000b). The authors suggested
therefore that the amino acid composition of nests may reflect phylogenetical rela-
tionships among wasp taxa. In addition, they also suggested that the evolution of

Nest materials in Polybia paulista
353
the chemical nature of the oral secretion, such as amino acid composition was also
important for social evolution in the family Vespidae.
The New World polistine wasps have developed greatly diverse societies, but
the kinds of nest material and their chemical nature have not yet been well studied.
In particular, no findings has been reported for the amino acid composition of the
nest protein of these wasps. In the present paper, we report the kind of nest materi-
al, nitrogen content and amino acid composition of the nests in a New World
swarm-founding polistine wasp, Polybia paulista.
MATERIALS AND METHODS
The wasps
Polybia paulista is one of the common swarm-founding social wasps in São Paulo state
(RICHARDS 1978). Colony size varies greatly (KUDÔ unpubl.), and is sometimes more than ten
thousand individuals (NOLL & ZUCCHI 2000). P. paulista colonies build ovoid nests that are
enclosed in a covering envelope with a single entrance hole. ITÔ et al. (1997) reported the
early developmental process of nests in this species, and NOLL & ZUCCHI (2000) have recently
made morphological analyses of this species.
Nest collection
Five colonies of Polybia (Myrapetra) paulista were collected on February 12-19, 1999 in
Ribeirão Preto (21˚11’S, 47˚48’W), southeastern Brazil. Three pre-emergence nests (nest 2, 3,
4) and two post-emergence nests (nest 1, 5) were used for analyses.
Observation of fine surface structure
Small fragments cut from the envelope and cell walls of nest 1 and 2 were observed
with a SEM (Nihon-denshi, JSN-T20), after coating with gold by an ion sputter (Nihon-
denshi, JFC-1100).
Elemental analysis
The nitrogen content of the upper part of envelope and cell walls at the first comb of
nest 1 were analyzed by a CHN-corder (Yanagimoto Seisakusho, Co. Ltd, Type MT-5) at the
Center for Instrumental Analysis, Ibaraki University.
Amino acid analysis
Small nest fragments, which were cut from envelopes of three nests (nest 3, 4, 5) and
cell walls of two nests (nest 3, 4), were weighed to 0.1 mg and hydrolyzed in 6N HCl at 110
˚C for 24 hr in sealed, evacuated tubes. Amino acids were then analyzed with an amino acid
analyzer, JEOL-300, which outputs an amount (in nmol) of each kind of amino acid. We
divided the total amount (nmol) of amino acids by the total sample weight (mg) in order to
estimate amount (nmol) of amino acids/mg of nest sample.

354
K. Kudô et alii
RESULTS
Fine surface structure
Nest materials
We first describe the nest materials used in nest 1. A SEM observation
revealed that minute vegetable chips (ca 1500 × 300 µm) were used as a major nest
material in all parts of the surface of both envelope and cell walls (Fig. 1). On the
surface of this plant material, there were several grooves (30-50 µm in width) that
were regularly oriented parallel to each other. From this appearance, such materi-
als are considered to be sieve tubes and/or vessels of plant tissues (Fig. 1A, D).
Numerous pits (ca 70 × 35 µm in major and minor diameters) (Fig. 1A) were also
seen on the surface of minute vegetable chips.
Fig. 1B shows that, in addition to the above plant tissues, Polybia paulista
also used plant hairs (ca 40 µm in width). Plant hairs are generally characterized in
the paper wasp nests, e.g., the nest material of Apoica, as thin and curled tissues
(YAMANE et al. in prep.).
Minor nest materials other than minute vegetable chips and plant hairs were
mud and/or inorganic particles (below to left margin in Fig. 1B). The appearance of
this minor nest material was very similar to that of eumenine and some stenogas-
trine wasps (KUDÔ et al. 1996), whose nests were completely composed of mud
(COWAN 1991, TURILLAZZI 1991).
The above description was also applicable to the other nest (nest 2), that is,
minute vegetable chips were a major component of the nest materials.
Oral secretion
Oral secretion was mainly used for binding plant tissues and other nest mate-
rials (Fig. 1C). It was found by a SEM observation that very small amounts of oral
secretion were used for nest construction, compared with that used in Polistes nests
(KUDÔ et al. 1998, YAMANE et al. 1998). In fact, the coating was only seen on the
upper surface of the envelope in nest 1 (Fig. 1D). Fig. 1E is a SEM micrograph
taken at a high magnification on the surface of the nest entrance in the envelope in
nest 2, showing the occurrence of the secretion coating.
Nitrogen content
An elemental analysis indicates that nitrogen (N) was contained in samples
from all the nests (n = 20). N content is a good index to indicate the amount of
oral secretion contained in the nests. In nest 1, mean ( SE) N content in the upper
parts of envelope, close to the first comb, was 2.01 0.04% (n = 10, range: 1.91-
2.14%), and that in the cell walls of the first comb was 1.71 0.04% (n = 10, range:
1.59-1.82%). There was a significant difference in N content between these nest
parts (Mann-Whitney U-test, P < 0.005). This supports the SEM observation that
the amount of oral secretion contained in the upper part of envelope was more
than that in the cell walls of nest 1. Further, it suggests that P. paulista applied dif-
ferent amounts of the secretion to different parts of the nest.

Nest materials in Polybia paulista
Amino acid composition of protein in the envelope and cell walls
355
The amino acid composition of protein in the envelope of three nests (nests 3,
4, 5) and the cell walls of two nests (nests 3, 4) was analyzed. Twenty-four amino
acids were detected from the five samples, of which the 15 dominant amino acids
are shown in Table 1. Twenty-two amino acids except for urea and L-α-aminoadipic
Fig. 1. — Scanning electron micrographs of: (A) the outer surface of the upper part of the envelope;
(B) the cell wall; (C) the cell wall; (D) the outer surface of the upper part of the envelope in nest 1;
and (E) the nest entrance on the envelope in nest 2. Scale bars = 50 µm in (A), (B) and (D), 30 µm
in (C), 25 µm in (E).

356
K. Kudô et alii
Table 1.
Amino acid composition (%) of protein from the envelope and the cell walls in three nests of Poly-
bia paulista.
Nest 3
Nest 4
Nest 5
Aminoacids
envelope
cell wall
envelope
cell wall
envelope
Ser
Gly
Ala
12.9
11.6
10.2
10.2
7.3
7.5
7.2
5.7
4.9
13.7
11.8
10.2
10.0
7.0
6.5
6.5
4.8
4.3
17.9
13.6
12.1
12.2
8.5
6.0
5.9
4.8
4.1
16.0
13.1
12.4
11.4
7.9
7.1
7.1
5.5
4.9
14.7
12.4
11.2
10.5
7.9
7.0
6.9
5.4
5.0
Val
Pro
Asp
Glu
Leu
Thr
H-Pro
Ile
5.2
3.3
4.5
2.8
1.7
3.0
5.1
3.6
1.4
3.2
Lys
3.1
2.6
2.2
3.0
2.5
Phe
Cit
2.5
2.3
2.0
2.2
1.7
1.0
2.2
1.5
2.2
3.2
Arg
2.1
1.6
1.7
2.6
2.0
acid were seen in all the samples analyzed. Seven amino acids, such as serine (15.0
0.9%, mean SE), glycine (12.5 0.4%), alanine (11.2 0.5%), valine (10.9
0.4%), proline (7.7 0.3%), aspartic acid (6.8 0.3%) and glutamic acid (6.7
0.2%), were major components, and these seven amino acids occupied approxi-
mately 70% of the protein. The proportions of these amino acids were similar for
the envelope and cell walls, suggesting that P. paulista use oral secretion with simi-
lar chemical components and composition in these two areas of the nest.
The mean amount of amino acids per mg in the above five samples was esti-
mated at 591.2 38.4 nmol. This value will be used for a comparison with those of
other known Polistes species in the discussion.
DISCUSSION
Nest materials
Polybia paulista uses three kinds of nest material, i.e., minute vegetable chips,
plant hairs and a small amount of mud and/or inorganic particles. According to
WENZEL (1991), wasps of genus Polybia utilize a great variety of nest materials. In
addition to the above nest materials, some Polybia species use long woody fibres
like Polistes (WENZEL 1991), but in the case of P. paulista, woody fibers were never
found in the nests.
We cannot conclude that P. paulista regularly collects mud and/or inorganic
particles in the field, because a possibility that such a material contaminated the
plant materials is not excluded. However, contamination with inorganic particles
has never been detected during SEM observations of nest material, which consists
only of long woody fibres, in two Japanese Polistes wasps, P. chinensis (KUDÔ et al.

Nest materials in Polybia paulista
357
1998) and P. riparius (YAMANE et al. 1998). In the absence of the accidental mixture
of the inorganic substance in these Polistes nests, it is possible that P. paulista col-
lected mud and/or inorganic particles as one of its nest materials. Two subgenera of
Polybia, Pedothoeca and Furnariana, rely solely on mud for their nest building
(RICHARDS 1978, WENZEL 1991). P. paulista belongs to another subgenus, Myrapetra,
but there is still a possibility that this species partially depends on some inorganic
materials for its nest building.
Factors determining the amount of oral secretion used for nest building
We found from a SEM observation that P. paulista uses only a small amount
of oral secretion for the construction and maintenance of their nests. This was sup-
ported by the low nitrogen contents (1.59-2.14%) of the envelopes and cell walls.
These values were much smaller than those for other known polistine species
(Polistes fuscatus: 6.6%, MCGOVERN et al. 1988; P. annularis: 2.8%, ESPELIE & HIM-
MELSBACH 1990; P. metricus: 1.4-8.0%, SINGER et al. 1992; P. chinensis: 1.3-4.2%,
KUDÔ unpubl.; Parapolybia indica: 6.4%, KUDÔ unpubl.; Ropalidia romandi: 2.7-
7.6%, KUDÔ unpubl.; R. montana: 2.9-5.0%, KUDÔ unpubl.), but nearly comparable
to those of Vespa species (KUDÔ unpubl.) (V. analis: 1.1-2.0%, V. simillima: 0.9-
2.0%, V. crabro: 2.5%). In general, the nitrogen content in most Polistes nests varies
considerably (ESPELIE & HIMMELSBACH 1990, SINGER et al. 1992), but the variation is
not very high in the nests of P. paulista and Vespa species. Large variances in nitro-
gen content between different parts of the nest in most polistine wasps may be due
to differences in the frequency of licking different areas with oral secretion. In
other words, these wasps would smear different nest parts with different amounts
of the secretion.
In P. paulista, the mean amount of amino acids/mg was estimated at 591.2
38.4 nmol, and this value was much smaller than those of two Japanese consub-
generic Polistes species, P. chinensis (1159.5
184.7 nmol, n = 9, KUDÔ unpubl.
data, Mann-Whitney U-test, P < 0.05) and P. riparius (1295.5 127.0 nmol, n = 5,
KUDÔ et al. 2000a, U-test, P < 0.01). This also supports that the amount of oral
secretion contained in P. paulista nests was much smaller.
P. paulista builds an envelope to cover their nest combs, of which the first
one is directly attached to the substrate (e.g., under a leaf of a palm tree) without
making pedicels (ITÔ et al. 1997). As stated above, although P. paulista uses a very
small amount of oral secretion, it builds large nests covered by an envelope. How
can this species build large nests using only a small amount of the secretion that
functions as an adhesive? WENZEL (1991) suggested that a notable feature of the
nests of Polybia and its nearest relatives, such as Brachygastra and Epipona is the
lack of a conspicuous matrix of secretion binding together the vegetable chip mate-
rial of the carton (Charterginus is exceptional). He considered that the reduced role
of secretion in the genera allied to Polybia may be due in part to the division of
labour in nest construction (see JEANNE 1986 for Polybia occidentalis). In addition
to his interpretation, there is a possibility that wasps do not need to use ample
secretion to bind minute vegetable chips. Interestingly, there are several facts that
support this interpretation. One of the facts is that Vespa species, which primarily
collect minute vegetable chips for nest building (WENZEL 1991, 1998; KUDÔ pers.
obs.), use a small amount of oral secretion for building and maintaining the nests
(see nitrogen content; WENZEL 1991). In contrast, some New World swarm-founding
polistine wasps, such as Protonectarina and Brachygastra mellifica that use woody

358
K. Kudô et alii
fibres, use more secretion than their near relatives, such as Polybia, Brachygastra
and Epipona (WENZEL 1991).
There are, however, several cases that are inconsistent with our interpretation.
For example, the basic nest material of two Old World swarm-founding polistine
species, Ropalidia romandi and R. montana is minute vegetable chips (KUDÔ
unpubl.), but these species build large nests covered by an envelope with much
larger amounts of oral secretion (see nitrogen content for these species; YAMANE &
ITÔ 1994). Furthermore, according to YAMANE & ITÔ (1994), R. romandi changes the
amount of oral secretion used for nest maintenance in response to environmental
conditions, such as precipitation. This fact suggests that the amount of oral secre-
tion used is determined by environmental factors, but not by division of labour and
nest materials. Flexible behaviours in smearing the nest surface with oral secretion
in response to precipitation have also been reported in the field colonies of P. chi-
nensis (KUDÔ et al. 1998, KUDÔ 2000).
It is very important to know whether the amount of oral secretion used is
determined by division of labour, nest materials or environmental factors. Our cur-
rent information would lead to the following interpretation. P. paulista builds their
nests using primarily minute vegetable chips, and further this species may have a
distinct division of labour in nest construction as measured by other Polybia
species (JEANNE 1986). Therefore, this species does not use a large amount of the
secretion. In addition, since most of their nests were built under the eaves of artifi-
cial buildings (sheltered sites) (SIMÕES & MECHI 1983) and palm trees (ITÔ et al.
1997), the wasps do not need to smear the nest surface frequently with the secre-
tion for maintenance. It may be plausible that the amount of oral secretion used
for building itself may be primarily determined by a division of labour in nest con-
struction and nest materials. In addition, wasps may change the amount of secre-
tion used for maintenance in response to given environmental conditions, e.g. pre-
cipitation (YAMANE & ITÔ 1994, KUDÔ et al. 1998, KUDÔ 2000).
It is important to know how the different amount of the secretion and the dif-
ferent kind and dimension of the construction materials affect the mechanical
properties of the nests in social wasps. COLE et al. (2001) recently analyzed the
physical properties of nest paper in three species of vespine wasps, and concluded
that the physical properties of nest paper could result from the collection of pulp
from different sources. In P. paulista, also, the physical properties of the nest mate-
rials should be analyzed in future work, because the capability for building big and
resistant nests is considered to be very important for the evolution of larger and
more complex colonies.
Amino acid composition of protein in envelope and cell walls
In P. paulista nests, seven dominant amino acids, i.e., serine, glycine, alanine,
valine, proline, aspartic acid and glutamic acid, occupied nearly 70% of the protein.
The first five amino acids were also dominant in the nests of Polistes (P. annularis,
ESPELIE & HIMMELSBACH 1990; P. metricus, SINGER et al. 1992; P. riparius, KUDÔ et
al. 2000a; P. chinensis, KUDÔ et al. 2000b) and Dolichovespula maculata (MCGOVERN
et al. 1988), although their proportions are considerably different. For example, the
proportion of glycine in the nests of P. chinensis (KUDÔ et al. 2000b) and P. riparius
(KUDÔ et al. 2000a) is much greater than in that of P. paulista. These data are con-
sistent with a view that the amino acid composition of protein in the nests reflects
phylogenetical relationships among social wasp taxa (KUDÔ et al. 2000a, 2000b).

Nest materials in Polybia paulista
359
In P. paulista, amino acid composition of the protein was similar among nests
and nest parts (envelope and cell walls). This suggests P. paulista produce oral
secretion with similar amino acid components and compositions regardless of the
parts of the nests and nesting environments. KUDÔ et al. (2000b) also found in pre-
emergence nests of P. chinensis that amino acid composition of protein in nest
papers and the secretion membrane covering the petiole surface were very similar
among nests built at different localities and nesting environments. They concluded
that the foundresses produced a similar kind of the secretion for construction and
maintenance of their nests.
Proline is a dominant component of structural proteins (BURSELL 1981, CHEN
1985). In spite of the small amount of secretion used in P. paulista nests, a relative-
ly high proline content (about 7.7%) would increase the mechanical strength of the
nest paper (ESPELIE & HIMMELSBACH 1990, SINGER et al. 1992, KUDÔ et al. 2000b).
The large amounts of aspartic acid and glutamic acid detected are a characteristic
feature seen in D. maculata nests (MCGOVERN et al. 1988). According to WENZEL
(1991), the nest materials in Polybia and Dolichovespula are very similar. It would
be very interesting to know the relationships between the kinds of nest materials
and amino acid composition, especially in the New World polistine wasps that have
greatly diversified.
ACKNOWLEDGMENTS
We express cordial thanks to two anonymous referees for their constructive comments
on the draft, and H. Hanawa, the Center for Instrumental Analysis, Ibaraki University for
analyzing the CHN composition of the nest material. This study was supported in part by a
Grant-in-Aid for Basic Researches from the Japan Ministry of Education, Science and Culture
(No. 10041154 and 12575004) to Sô. Yamane and Research Fellowships of the Japan Society
for the Promotion of Science for Young Scientists to K. Kudô. This study was also supported
in part by a Grant-in-Aid for Basic Researches from the Japan Ministry of Education, Science
and Culture (No. 12304048). The collection of nest samples was permitted by Ofício 115/00
DIFAS/DIREC, Licença 074/00, Favorecido R. Zucchi e outros.
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