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A phylogenetic analysis of Synoeca De Saussure, 1852, a neotropical genus of social wasps (hymenoptera: vespidae: epiponini).

Abstract.--Characters of adult morphology, male genitalia, and nest architecture are combined in a cladistic analysis of the neotropical paper wasp genus Synoeca. The single resulting cladogram supports the monophyly of genus. The species are related as follows: S. chalibea + (S. virginea + (S. septentrionalis + (S. surinama + S. cyanea))). A new identification key is also presented.

Key words. Vespidae, Epiponini, Synoeca, cladistics.

**********

Synoeca is a small genus in the paper wasp tribe Epiponini, comprising five species (Synoeca chalibea de Saussure, S. virginea F., S. surinama L., S. septentrionalis Richards and S. cyanea F.). Individuals are usually medium-sized and some species have metallic colors. The genus has a wide distribution in Central and South America, extending from Mexico to Argentina (Richards, 1978). The nests are familiar structures throughout the range: arboreal, usually on a broad slanting surface and with a single sessile comb attached directly to the tree trunk (Wenzel, 1998). When disturbed, warning behavior consisting of drumming or tapping inside the nest is audible (Overal, 1982; O'Donnell et al., 1997), and their aggressive defense of the obvious nests has led to common names such as "seven mile jep" or "guitarron."

Carpenter (1991) presented a phylogenetic analysis of genera of Polistinae, with Synoeca placed as sister-group of Clypearia (= Occipitalia; synonymized with Clypearia by Carpenter et al., 1996), Asteloeca and Metapolybia, but the interrelationships among the latter genera were unresolved. Wenzel (1993) and Wenzel and Carpenter

(1994) subsequently presented analyses with different resolutions of relationships among the four genera (see discussion below).

Recent behavioral studies have been published on Synoeca (Schremmer, 1973; Bermudez, 1980a, 1981a & b, 1982; West-Eberhard, 1981; Thiago et al., 2005), including detailed treatments of caste differentiation (Noda et al., 2003; Noll et al., 2004) and nest architecture (Bermudez, 1980b; Overal, 1982; Schremmer, 1986; Wenzel, 1991, 1998), but there has been no investigation of species relationships. In the present work a phylogenetic analysis of Synoeca using characters of external morphology, male genitalia and nest architecture is given. A new illustrated identification key based on this phylogenetic analysis combined with data from Richards' (1978) key is also presented.

MATERIAL AND METHODS

A total of 86 specimens were examined in this work, from the American Museum of Natural History, New York, USA (AMNH), and Faculdade de Filosofia, Ciencias e Letras de Ribeirao Preto da Universidade de Sao Paulo, Ribeirao Preto, Brazil (FFCLRP-USP) (see numbers and localities for species in Appendix 1).

Twenty-five external morphological characters, one from male genitalia, and twelve nest characters were scored in a data matrix for phylogenetic reconstruction (Tables 1 and 2). The morphological characters were examined on pinned specimens, male genitalia were examined by dissection, while the nest characters were taken from the literature (Wenzel, 1993, 1998). Some of these characters are illustrated in Figs. 1-8. The figures of the male genitalia of S. virginea (from Brazil, Para state, Fig. 7), and S. surinama (locality not specified, Fig. 8) were originally part of a parcel of unpublished figures of polistine genitalia by the late J. van der Vecht given to J. M. Carpenter.

Multistate characters were treated as additive, except characters 3, 18-19, 23, 26 and 33. Cladistic analysis was undertaken using the program TNT (Tree analysis using New Technology; Goloboff et. al., 2003). Outgroup rooting (Nixon and Carpenter, 1993) was implemented with the following taxa: Charterginus nevermanni Bequaert, Epipona guerini (de Saussure), Metapolybia docilis Richards, Asteloeca traili (Cameron), and Clypearia weyrauchi Richards.

RESULTS

Analysis of the matrix by implicit enumeration resulted in a single cladogram of length 67, consistency index 0.83 and retention index 0.81 (Fig. 9). The same cladogram resulted from implied weighting (default concavity). Synoeca is supported as a monophyletic genus, with relationships among the species: S. chalibea + (S. virginea + (S. septentrionalis + (S. surinama + S. cyanea))).

DISCUSSION

Carpenter's (1991) analysis of generic relationships in Polistinae, based on morphological characters, established a monophyletic group comprising Asteloeca, Clypearia (= Occipitalia) and Metapolybia, based on the anterior pronotal carina raised (in our matrix as character 10), but did not resolve the interrelationships among these three taxa. In our analysis other morphological characters supporting this group include scutellum compressed (character 17, state 1), first metasomal tergum and sternum without numerous erect hairs outstanding above dense pubescence (characters 20, state l, and 21, state 1), and nest envelope reinforcement by secretion (character 30, state 1). The sister-group of these three genera is Synoeca, also proposed by Carpenter (1991), here based on nest characters: comb without pulp foundation (character 29, state 1), envelope not wide at bottom (character 31, state 1), entrance a short collar-like structure (character 32, state 1) and envelope closes during cell outlining (character 38, state 0). Wenzel's (1993) analysis of nest architecture also recognized a lineage comprising those four genera. Although the characters supporting this clade were not detailed, it was based on three characters: comb built on bark without pulp foundation, material of coarse chips, and envelope reinforcement by

secretion (Carpenter et al., 1996). In fact, Synoeca has envelope reinforcement by blots, not by secretion (Wenzel, 1993: table 1), so this character supports only the clade comprising Metapolybia, Asteloeca and Clypearia. Nest material composed of short chips is a feature shared by those four genera, but it is also present in Epipona, hence is not a synapomorphy on Fig. 9.

[FIGURE 1 OMITTED]

[FIGURE 2 OMITTED]

[FIGURE 3 OMITTED]

[FIGURE 4 OMITTED]

[FIGURE 5 OMITTED]

[FIGURE 6 OMITTED]

[FIGURE 7 OMITTED]

[FIGURE 8 OMITTED]

[FIGURE 9 OMITTED]

Wenzel and Carpenter (1994) combined the data matrices from Carpenter (1991) and Wenzel (1993), and added larval characters provided by J. Kojima. Their consensus tree resolved Asteloeca + Metapolybia as sister-group of Clypearia, and Synoeca as sister-group of these three genera. In all of these studies Synoeca is the most basal genus, except for Wenzel (1993), which had the topology for these four genera as: Clypearia (=Occipitalia) + (Asteloeca + (Metapolybia + Synoeca)). The most basal placement of Synoeca is confirmed in this study.

Synoeca is here supported by: vertex projecting (character 1, state 0; Figs. 1-2), which is also a diagnostic feature for Synoeca in Richards (1978); and nest entrance formed by gap in construction (character 34, state 1; homoplastic in Fig. 9 because it is also found in Metapolybia). Several other features are shared by all the Synoeca species, including fore coxae rounded (character 14, state 1; also mentioned by Carpenter, 1991), propodeum with a slight concavity above propodeal orifice (character 18, state 1) and the first metasomal segment filiform basally, widening gradually after spiracles (character 19, state 2). Although some of the states shown by Synoeca are unique, because of variation in its sister-group, the optimization of these characters is ambiguous, hence they are not plotted on Fig. 9. Another character, the prestigma (character 25), is also ambiguous. According to Richard's (1978) key, the state would be as long as wide (state 0) for S. septentrionalis and S. cyanea. However, we have observed that these species have the prestigma longer than wide, as in the other species of Synoeca. In any case, generic monophyly is well established.

In Richards' (1978) key, the punctation on the pronotum, scutum, scutellum, and propodeum are important diagnostic features to recognize Synoeca species, and some of these features are synapomorphic in our cladogram. Concerning the propodeal punctation (character 16), Synoeca might be separated into two groups: (1) S. chalibea and S. virginea, with propodeal punctation present and dense (state 0, Fig. 3); and (2) S. surinama, S. cyanea and S. septentrionalis, with propodeal punctation reduced in part dorsally and laterally (state 2, Fig. 4). As already shown by the key of Richards (1978), species of these two groups have the following diagnostic differences: (character 4, clypeal-eye contact) clypeus touching the eyes vs. clypeus narrowly separated from the eyes; (character 5, malar space) malar space shorter than pedicel (Fig. 1) vs. about as long as pedicel (Fig. 2); and (character 26, wing color) wings yellowish to brownish, darker anteriorly vs. infuscate throughout. In addition, these two groups are differentiated by the color of the body as a whole (yellowish vs. bluish to blackish), a character not included in our matrix.

Synoeca chalibea is the only species within Synoeca that has dense punctation on the pronotum, scutum and propodeum, sharing these features with the most outgroups, placing this species as sister-group to the remainder of the genus. Thus, although S. chalibea and S. virginea share several features, the sparse punctation on the pronotum and scutum (Fig. 6) in S. virginea groups it phylogenetically with S. surinama, S. cyanea and S. septentrionalis. There has evidently been a reduction trend in body punctation in the genus, because the loss of punctures found in S. surinama is derived relative to the other species. Finally, it is two other reduction characters, first metasomal tergum and sternum without numerous erect outstanding hairs (characters 20-21), that resolves S. surinama and S. cyanea as sister-groups. However, this is homoplastic, as the condition is found in several outgroups as well. Richards (1978) mentioned this condition for the tergum in those two species at two places in his key, characterizing it as "First gastral tergite with few outstanding hairs" for S. surinama, and "fewer outstanding hairs" for S. cyanea. These species show a dense pubescence on the anterior region of the tergum, as do S. septentrionalis, S. chalibea and S. virginea, but not outstanding hairs. We also have seen that the outstanding hairs on sternum I show the same pattern of reduction as that on tergum I, being another shared feature for the clade S. surinama + S. cyanea.

A couple of other characters deserve special comment. Hitherto, the male genitalia in Synoeca had not been properly described, and the unique genitalia character included in our matrix (character 24, digitus of male genitalia), supports the grouping of S. surinama, S. cyanea and S. septentrionalis. These species shared the posterodorsal lobe sharply pointed (state 1), while S. chalibea and S. virginea share with the outgroups the lobe rounded (Figs. 7-8, and cf. du Buysson, 1906: pl. 15 figs. 8-9). Conditions approaching these two states have been seen in species of other genera not included here (Carpenter and Mateus, 2004; Andena et. al., 2007), and although not identical, several independent origins of similar states in Epiponini seems probable.

The structure called the thyridium is a small area of differentiated cuticle located at on each side of the base of the second metasomal tergum, connected with the front margin by a narrow Stalk (Richards, 1978). Its shape is variable in Synoeca, but Richards (1978: 178) stated: "Its shape, though variable, is of some taxonomic importance." It is included in our matrix as characters 22 and 23, and in fact is uninformative. In S. virginea and S. cyanea this structure is coded as elongate, widening gradually (character 23, state 2), while in the remaining species of Synoeca the shape of thyridium is subcircular (state 1). We have seen variation in the shape of thyridium in all species within Synoeca, but these variants are not enough to be coded as different states. States different from either of these are found in each of the outgroups. Within Synoeca, therefore, the evolution of this character is ambiguous.

IDENTIFICATION KEY FOR THE SPECIES OF SYNOECA DE SAUSSURE
1. Malar space shorter than pedicel (Fig. 1);
propodeum with fine punctation throughout
(Fig. 3); wings yellowish to brownish, darker
anteriorly, body yellowish brown to bluish or
greenish                                                            2
Malar space about as long as pedicel (Fig. 2);
propodeum with punctation sparse or evanescent
at borders (Fig. 4); wings infuscate
throughout, body bluish to blackish                                 3

2. Pronotum, scutum and scutellum with fine,
dense punctation (Fig. 5).                                          3
                          chalibea de Saussure
Pronotum, scutum and scutellum with sparse
punctation (Fig. 6), sometimes, impunctate.
virginea (F.)

3. Pronotum, scutum and scutellum impunctate;
propodeum with sparse punctation; clypeus
black, mandibles, malar space and lower gena
black to brown-marked.                                   surinama (L.)
Pronotum scutum and scutellum with sparse
punctation; propodeum with punctation denser;
clypeus, mandibles, malar space and lower
gena reddish-marked.                                                 4

4. First metasomal tergum, and sternum lacking
erect hairs outstanding above dense pubescence
basally; clypeus reddish.                                   cyanea (F.)
First metasomal tergum and sternum with
numerous erect hairs basally; clypeus with a
more or less dark triangular area dorsally.    septentrionalis Richards


Received and accepted January 21, 2008.

APPENDIX 1.

LOCALITIES FOR SPECIMENS EXAMINED.

S. chalibea: Brazil [Obidos, Amazonas State, 19 (AMNH); Rio Branco, Acre State, 2 [female][female] (FFCLRP-USP); Nova Mutum, Mato Grosso State, 8 99 (FFCLRP-USP)]; Peru [Col. Perene, Pasco, 1[female]1[male] (AMNH); Rio Santiago, Loreto, 19 (AMNH); Satipo, Junin, 19 (AMNH); El Campeziento, 1[female] (AMNH)]; Costa Rica [Golfito, Puntarenas, 19 (AMNH)].

S. cyanea: Argentina [Misiones, 19 (AMNH)]; Brazil [Nova Teutonia, Santa Catarina State, 1[female]1[male] (AMNH); Pedregulho, Sao Paulo State, 3[female][female] (FFCLRP-USP); Sta Rosa de Viterbo, Sao Paulo State, 19 (FFCLRP-USP); Osorio, Rio Grande do Sul State, 399 (FFCLRP-USP)]; Paraguay [Villa Rica, Guaira, 1[female]1[male] (AMNH)].

S. septentrionalis: Colombia [Rio Frio, Magdalena, 19 (AMNH)]; Panama [Coanguinola Dist., 1[male] (AMNH)]; Guatemala [Sta Emilia, Pochuta, 1[male] (AMNH)], Honduras [Zamorano, 19 (AMNH)]; Mexico [Muste, Chiapas, 299 (AMNH)].

S. surinama: Bolivia [El Porvenir, Beni, 1[female] (AMNH)]; Brazil [Uypiranga, Amazonas State, 1[male] (AMNH); Obidos, Para State, 19 (AMNH); Araguaia River, Goias State, 299 (FFCLRP-USP); Nova Mutum, Mato Grosso State, 19 (FFCLRP-USP); Nova Xavantina, Mato Grosso State, 19 (FFCLRP-USP), Lizarda, Tocantins State, 23[female][female] (FFCLRP-USP)]; Suriname [Paramaribo, 19 (AMNH); Raleighvallen-Voltzberg, 1[female] (AMNH)]; Trinidad [St. Augustine, 1[female] (AMNH)].

S. virginea: Brazil [locality not specified, 1[female] (AMNH); Rio Branco, Acre State, 1[female] (FFCLRP-USP); Nova Mutum, Mato Grosso State, 1[female] (FFCLRP-USP)]; Peru-Brazil frontier, 1[female] (AMNH); Ecuador [Coca, Orellana, 19 (AMNH); Limoncocha, Napo, 89 (AMNH)]; Colombia [La Chorerra, Amazonas, 1[female]2[male][male] (AMNH)]; Suriname [Raleighvallen-Voltzberg, 19 (AMNH)].

ACKNOWLEDGMENTS

We thank Kurt M. Pickett and Ward Wheeler for assistance during S. R. Andena's visit to the American Museum of Natural History; and Ronaldo Zucchi (FFCLRP-USP) for assistance in Ribeirao Preto. This work was supported by grants from

Fundacao de Amparo a Pesquisa do Estado de Sao Paulo (04/01117-0; 01/02491-4), CNPQ (Conselho Nacional de Desenvolvimento Cientifico e Tecnologico), and a Collection Study Grant (American Museum of Natural History) to S. R. Andena.

LITERATURE CITED

Andena, S. R., F. B. Noll, J. M. Carpenter and R. Zucchi. 2007. Phylogenetic analysis of the neotropical Pseudopolybia de Saussure, 1863, with description of the male genitalia of Pseudopolybia vespiceps (Hymenoptera: Vespidae, Epiponini).

American Museum Novitates 3586:1-11.

Bermudez, E. G. C. 1980a. Reproducao e dinamica de populacao em Synoeca surinama (Hymenoptera: Vespidae). Acta Amazonica 10(3): 679-690.

Bermudez, E. G. C. 1980b. Orientacao, arquitetura e construcao dos ninhos por Synoeca surinama L. (hymenoptera: Vespidae). Acta Amazonica 10(4): 883-896.

Bermudez, E. G. C. 1981a. Comportamento e causas da migracao de Synoeca surinama L. (Hymenoptera: Vespidae) na amazonia Brasileira. Revista Brasileira de Biologia 26(1): 71-74.

Bermudez, E. G. C. 1981b. Alarme e defesa no ninho de Synoeca surinama (Hymenoptera: Vespidae). Acta Amazonica 11(2): 377-382.

Bermudez, E. G. C. 1982. Evidencias sobre o comportamenLo de copula dos machos de Synoeca surinama L. (Hymenoptera: Vespidae) fora do ninho. Acta Amazonica 12(3): 665-666.

Buysson, R. du. 1906. Monographie de Vespides appartenant aux genres Apoica et Synoeca. Annales de la Societe Entomologique de France 75: 333-362.

Carpenter, J. M. 1991. Phylogenetic relationships and the origin of social behavior in the Vespidae. In: K. G. Ross and R. W. Matthew (eds.), The social biology of wasps, 7-32. Cornell University Press, Ithaca, NY.

Carpenter, J. M. and S. Mateus. 2004. Males of Nectarinella Bequaert (Hymenoptera, Vespidae; Polistinae). Revista Brasileira de Entomologia 48(3): 297-302.

Carpenter, J. M., J. W. Wenzel and J. Kojima. 1996. Synonymy of the genus Occipitalia Richards, 1978, with Clypearia de Saussure, 1854 (Hymenoptera: Vespidae; Polistinae, Epiponini). Journal of Hymenoptera Research 5: 157-165.

Goloboff, P., S. Farris and K. Nixon. 2003. T. N. T.: Tree analysis using New Technology. Published by the authors, Tucuman, Argentina.

Nixon, K. C. and J. M. Carpenter. 1993. On outgroups. Cladistics 9: 413-426. Noda, S. C. M., S. N. Shima and F. B. Noll. 2003.

Morphological and physiological caste differences in Synoeca cyanea (Hymenoptera, Vespidae, Epiponini) according to the ontogenetic development of the colonies. Sociobiology 41: 547-570. Noll, F. B., J. W. Wenzel and R. Zucchi. 2004. Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini). American Museum Novitates 3467: 1-24.

O'Donnell, S., J. H. Hunt and R. L. Jeanne. 1997. Gaster-flagging during colony defense in neotropical swarm-founding wasps (Hymenoptera: Vespidae, Epiponini). Journal of the Kansas Entomological Society 70(3): 175-180.

Overal, W. L. 1982. Acoustical behavior and variable nest architecture in Synoeca virginea (Hymenoptera: Vespidae). Journal of the Georgia Entomological Society 17(1): 1-4.

Richards, O. W. 1978. The social wasps of the Americas excluding the Vespinae. British Museum (Natural History), London.

Schremmer, F. 1973. Synoeca cyanea (Vespidae). Alarmverhalten und Nestreparatur (Freilandaufnahmen). In: G. Wolf (ed.), Encyclopaedia Cinematographica, Film E 1886. Institut fur Wissenschaftlichen Film, Gottingen.

Schremmer, F. 1986. Die Bautenvielfalt der sozialen Wespen. Oko-L 8(2-3): 49-59. Thiago, E., C. Ribeiro Junior, D. L. Guimaraes and F. Prezoto. 2005. Foraging activity and nesting of swarm-founding wasp Synoeca cyanea (Hymenoptera: Vespidae, Epiponini). Sociobiology 46(2): 317-327.

Wenzel, J. W. 1991. Evolution of nest architecture in social vespids. In: K. G. Ross and R. W. Matthews (eds.), The Social Biology of Wasps, 480-519. Cornell University Press, Ithaca, NY.

Wenzel, J. W. 1993. Application of the biogenetic law to behavioral ontogeny: a test using nest architecture in paper wasps. Journal of Evolutionary Biology 6: 229-247.

Wenzel, J. W. 1998. A generic key to the nests of hornets, yellowjackets, and paper wasps worldwide (Vespidae: Vespinae, Polistinae). American Museum Novitates 3224: 1-39.

Wenzel, J. W. and J. M. Carpenter. 1994. Comparing methods: adaptative traits and tests of adaptation.

In: P. Eggleton and R. Vane-Wright (eds.), Phylogenetics and Ecology, 79-101. Academic Press, London.

West-Eberhard, M. J. 1981. Intragroup selection and the evolution of insect societies. In: R. D. Alexander and D. W. Tinkle (eds.), Natural Selection and Social Behavior: Recent Research and New Theory, 3-17. Chiron, New York.

SERGIO R. ANDENA (1) *, JAMES M. CARPENTER (2) AND FERNANDO B. NOLL (3)

(1) FFCLRP-USP. Depto. Biologia. Av. Bandeirantes, 3900, Ribeirao Preto-SP, CEP 14040-901, Brazil. e-mail: sandena@usp.br

(2) Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA. e-mail: carpente@amnh.org

(3) Departamento de Zoologia e Botanica; Instituto de Biociencias, Letras e Ciencias Exatas, UNESP; Rua Cristovao Colombo, 2265; 15054-000, Silo Jose do Rio Preto, SP; Brazil. e-mail: noll@ibilce.unesp.

* Present address: Department of Biology, University of Vermont, 120 A Marsh Life Science Building, Burlington, VT 05405, USA. E-mail: sergio.andena@uvm.edu
Table 1. Character list for Synoeca.

Character                             States

1.          Vertex: projecting = 0; rounded = 1
2.          Shape of clypeus: longer than wide = 0; at least as long
              as wide = 1
3.          Apex of clypeus: rounded-truncate = 0; rounded = 1;
              emarginate = 2; truncate = 3
4.          Clypeal eye-contact: clypeus touching the eyes = 0; clypeus
              narrowly separated from the eyes = 1
5.          Malar space: shorter than pedicel (Fig. 1) = 0; about as
              long as pedicel = 1 (Fig. 2)
6.          Gena:: narrower than eyes = 0; equal to or wider than
              eyes = 1 (Figs. 1-2)
7.          Occipital carina: present = 0; absent = 1
8.          Mandibular edge: rounded = 0; raised into flange = 1
9.          Pronotal punctation: dense, coarse = 0; dense, fine = 1
              (Fig. 5); evanescent or absent = 2 (Fig. 6)
10.         Anterior pronotal carina: short, blunt = 0; raised = 1
11.         Dorsal pronotal carina: present = 0; absent = 1
12.         Pronotal fovea: present = 0; absent = 1
13.         Pretegular carina: absent = 0; present = 1
14.         Fore coare: laterally strongly produced = 0; rounded = 1
15.         Scutal punctation: dense, coarse = 0; fine = 1 (Fig. 5);
              evanescent or absent = 2 (Fig. 6)
16.         Propodeal punctation: present = 0 (Fig. 3); rugose = 1;
              reduced in part dorsally and laterally = 2 (Fig. 4)
17.         Scutellum: rounded = 0; compressed = 1
18.         Propodeal concavity: shallow, broad = 0; deep, low = 1;
              with a slight concavity above propodeal orifice = 2;
              narrow, not reaching metanotum = 3; shallow, narrow = 4;
              deep, narrow = 5
19.         First metasomal segment: petiolate, with wide part
              posteriorly = 0; filiform = 1; filiform basally,
              widening gradually after spiracles = 2; filiform basally,
              widening abruptly after spiracles = 3; filiform basally,
              little widened after spiracles = 4
20.         First metasomal tergum:: with numerous erect hairs
              outstanding above dense pubescence basally = 0; without
              numerous erect hairs outstanding above dense pubescence
              basally = 1.
21.         Sternum P with numerous erect hair = 0; without hairs, or
              reduced = 1
22.         Position of thyridium: basal = 0; distant from the anterior
              edge of second metastomal tergum = 1
23.         Thyridium shape: widely transverse = 0; subcircular = 1;
              elongate, widening gradually = 2; short, linear = 3;
              subpyriform = 4.
24.         Digitus of male genitalia: posterodorsal lobe rounded = 0
              (Fig. 7); posterodorsal lobe sharply pointed = 1 (Fig. 8)
25.         Prestigma: as long as wide = 0; longer than wide = 1
26.         Wing color: hyaline, infuscate anteriorly = 0; yellowish
              to brownish, darker anteriorly = 1; infuscate
              throughout = 2; hyaline = 3
27.         Nest material. long fibers = 0; short chips = 1
28.         Comb foundation: sessile, resembling petiole = 0;
              sessile = 1
29.         Comb with pulp foundation: present = 0; absent = 1
30.         Envelope reinforcement: by blots = 0; by secretion = 1
31.         Envelope breadth: widest at bottom = 0; not wider at
              bottom = 1
32.         Entrance: simple hole = 0; short collar-like structure = 1
33.         Entrance position: dorsal = 0; ventral = 1; central or
              toward periphery of envelope = 2
34.         Entrance construction: formed by remaining gap in
              construction = 0; formed as separate structure away from
              last gap in advancing edge = 1
35.         Secondary combs: absent or contiguous with primary
              comb = 0; present, on envelope = 1
36.         Secondary envelopes: absent = 0; present = 1
37.         Comb expansion: gradual = 0; successive blocks = 1
38.         Cells: most cells laid before envelope closes = 0;
              envelope closes during cell outlining = 1

Table 2. Matrix for species of Synoeca. The question mark (?) denotes
not seen.

                          1   2   3   4   5   6   7   8   9   10   11

Charterginus nevermanni   1   0   0   0   0   0   0   0   0    0    0
Epipona guerini           1   1   2   0   0   1   1   0   1    0    1
Synoeca surinama          0   1   1   1   1   1   1   0   2    0    1
Synoeca cyanea            0   1   1   1   1   1   1   0   2    0    1
Synoeca septentrionalis   0   1   1   1   1   1   1   0   2    0    1
Synoeca chalibea          0   1   1   0   0   1   1   0   1    0    1
Synoeca virginea          0   1   1   0   0   1   1   0   2    0    1
Asteloeca traili          1   1   1   0   0   1   1   1   2    1    1
Metapolybia docilis       1   1   1   0   0   1   1   1   1    1    1
Clypearia weyrauchi       1   0   3   0   0   0   1   0   1    1    1

                          12   13   14   15   16   17   18   19   20

Charterginus nevermanni    0    0    0    0    0    0    0    0    0
Epipona guerini            1    1    0    1    1    0    1    1    0
Synoeca surinama           1    1    1    2    2    0    2    2    1
Synoeca cyanea             1    1    1    2    2    0    2    2    1
Synoeca septentrionalis    1    1    1    2    2    0    2    2    0
Synoeca chalibea           1    1    1    1    0    0    2    2    0
Synoeca virginea           1    1    1    2    0    0    2    2    0
Asteloeca traili           1    1    0    2    0    1    3    3    1
Metapolybia docilis        1    1    0    1    0    1    4    4    1
Clypearia weyrauchi        1    1    1    1    0    1    5    3    1

                          21   22   23   24   25   26   27   28   29

Charterginus nevermanni    0    0    2    0    0    0    0    0    0
Epipona guerini            0    1    1    0    1    0    1    1    0
Synoeca surinama           1    1    1    1    1    2    1    1    1
Synoeca cyanea             1    1    2    1    1    2    1    1    1
Synoeca septentrionalis    0    1    1    1    1    2    1    1    1
Synoeca chalibea           0    1    1    0    1    1    1    1    1
Synoeca virginea           0    1    2    0    1    1    1    1    1
Asteloeca traili           1    1    3    0    0    3    1    1    1
Metapolybia docilis        1    1    3    0    0    3    1    1    1
Clypearia weyrauchi        1    1    4    ?    0    3    1    1    1

                          30   31   32   33   34   35   36   37   38

Charterginus nevermanni    0    0    0    0    0    0    0    0    1
Epipona guerini            0    0    0    1    0    1    1    1    1
Synoeca surinama           0    1    1    2    1    0    0    0    0
Synoeca cyanea             0    1    1    2    1    0    0    0    0
Synoeca septentrionalis    0    1    1    2    1    0    0    0    0
Synoeca chalibea           0    1    1    2    1    0    0    0    0
Synoeca virginea           0    1    1    2    1    0    0    0    0
Asteloeca traili           1    1    1    2    0    0    0    0    0
Metapolybia docilis        1    1    1    2    1    0    0    0    0
Clypearia weyrauchi        1    1    1    2    0    0    0    0    0
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Author:Andena, Sergio R.; Carpenter, James M.; Noll, Fernando B.
Publication:Entomologica Americana
Article Type:Report
Geographic Code:1USA
Date:Jan 1, 2009
Words:4135
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