The paper wasp genus Marimbonda Richards, 1978, is synonymized with Leipomeles Möbius, 1856, n.syn. A key to all the currently recognized paper wasp genera in the Western Hemisphere is provided.
INTRODUCTION
The paper wasp genus Marimbonda was described by Richards (1978) for two species: the type species M. albogrisea Richards, 1978, and M. pusilla (Ducke, 1904). The recognition of this new genus was due primarily to the nest architecture of the new species M. albogrisea. The single nest collected was astelocyttarus (Richards and Richards, 1951: 6): the comb lacking any petiole, with the cells constructed on a flat surface and an envelope covering the comb. Richards (1978: 10) keyed the new genus out with those genera also bearing a stout, curved bristle on the third segment of the labial palpus (figs. 1–2), and among these the genera sharing reduction in the palpal formula to five-segmented maxillary palpi and three-segmented labial palpi (fig. 1). However, his dendrogram (Richards, 1978: fig. 40) showed Marimbonda as intermediate between these genera and a group clade including other genera with astelocyttarus nest architecture. Richards (1978: 201) stated: “The genus whose relationships are shown in the dendrogram … and in the key … does not seem to be at all allied to the others which make astelocyttarus nests (Synoeca, Metapolybia, and Clypearia) and while it should probably be placed somewhere near the others which have the curved bristle on the labial palpus, it does not seem to be close to any of them.”
The other genera with the bristle on the third labial palpomere are Pseudopolybia, Parachartergus, Chartergellus, Nectarinella, and Leipomeles. In previous cladistic analyses of paper wasp genera (Carpenter, 1991; Wenzel and Carpenter, 1994) these genera were shown to form a monophyletic group including Marimbonda; that bristle is a synapomorphy. Within this clade, Marimbonda and Leipomeles are sister-groups. A morphological synapomorphy is the similar form of the first metasomal segment (conically petiolate; figs. 3–4), while a possible synapomorphy in nest architecture is a central furrow built in the lines of construction of the envelope.
In the present work, Marimbonda and Leipomeles are synonymized. Although it is possible to adduce morphological apomorphies to support each genus as monophyletic (e.g., Leipomeles has the first metasomal segment forming a longer, relatively narrower petiole, fig. 4; Marimbonda has the hindwing with vein Cu1 as long as cu-a, fig. 5, vs. Cu1 shorter than cu-a in Leipomeles, fig. 6), these are all minor features, relative both to the features shared by these genera and features defining other paper wasp genera. More significantly, Marimbonda was recognized as a genus in the first place because of its nest architecture, but its astelocyttarus design is less important than other features of the nest that it shares with Leipomeles (Wenzel and Carpenter, 1994). The astelocyttarus nest obscured the relationships of Marimbonda when that genus was described—in fact was misleading, as seen from Richards' placement of the genus on his dendrogram. Moreover, as will be shown, astelocyttarus nests do not in fact separate Marimbonda from Leipomeles.
After discussion of taxonomic history and further explication of the rationale for the synonymy, a new key to the paper wasp genera is provided, which will cover all the taxa found in the New World.
TAXONOMIC HISTORY
The present generic classification of New World Polistinae owes its basic form to the work of Adolpho Ducke. A century ago, working at the Museu Goeldi in Belem, he revised the classification then prevailing, and his views were especially informed by his own extensive field work in Amazonian Brazil. Prior to Ducke the classification used was basically that in the last worldwide monograph (de Saussure, 1853–1858), which itself was a refinement of Lepeletier's (1836) classification, who began the recognition of numerous genera for Polistinae. Polistinae were separated from other Vespidae (as the Linnaean genus Vespa) by Latreille (1802), who described two genera, Polistes and Epipona. Between the time of Latreille and Lepeletier just a few isolated genera were described (Kirby, 1826: Cyclostoma2; Guérin-Méneville, 1831: Ropalidia; Perty, 1833: Brachygastra), but Lepeletier described five genera: Agelaia, Apoica, Chartergus, Polybia, and Rhopalidia.3 Between the time of Lepeletier and de Saussure again just a couple of genera were added (White, 1841: Anthreneida,4 Myrapetra5). De Saussure (1852) added Synoeca and de Saussure (1853–1858) Raphigaster6 and Mischocyttarus. Subsequent to de Saussure's work a few more genera accumulated (Möbius, 1856: Leipomeles; Gribodo, 1892: Paraicaria7; H. von Ihering, 1896: Pseudopolybia8; Fox, 1898: Charterginus).
Working on the fauna of Pará, Ducke (1904) at first followed de Saussure's classification, adding just one genus, Megacanthopus.9 But shortly thereafter, Ducke (1905a) radically revised the generic classification of the South American Polistinae. De Saussure had recognized eight genera of Polistinae in the Neotropics; Ducke (1905a) recognized 18. In particular, he split up the genera Chartergus and Polybia, both quite heterogeneous in his 1904 study. With respect to Chartergus he added the genus Parachartergus, a genus described shortly before by R. von Ihering (1904), as well as the new genus Pseudochartergus. In addition to Polybia, he raised to generic rank Clypearia de Saussure, 1854, and Charterginus Fox, 1898, treated by him as subgenera of Polybia in 1904; recognized Leipomeles Möbius, 1856, treated by him as a synonym of Polybia in 1904; and added the new genera Synoecoides and Metapolybia.
Marimbonda pusilla was originally described in the genus Chartergus by Ducke (1904), from Belem and Oyapoc in Brazil. When Ducke (1905a) began to revise the generic classification, he transferred C. pusillus to Parachartergus. Möbius (1856) first used the palpal formula as a taxonomic character, when describing the genus Leipomeles, and R. von Ihering (1904: 129) cited this example in justifying his new genus. Parachartergus was described as monotypic, distinguished from Chartergus by labial palpi being three-segmented not four-segmented. Ducke (1905a, 1905b), however, argued that many species hitherto considered as Chartergus had the fourth labial palpomere rudimentary, and that therefore they could not be separated generically from Parachartergus. As defining characters of Parachartergus were cited the thick, long, curved bristle before the extremity of the labial palpus, and nests, where known, all stelocyttarus calyptodomous (terminology of de Sausure, 1853–1858, meaning petiole combs covered with an envelope and free of the latter). Interestingly, Ducke (1904: 357) had refused to accept Leipomeles as a genus, deriding the importance of the palpi as a generic character: “Mas como hoje já passou o tempo em que se baseava generos novos sobre pequenas differenças no apparelho buccal …”. Under his changed view (Ducke, 1905a), the five-segmented maxillary palpi was the key character state separating Leipomeles from Parachatergus, both of which had the bristle on the labial palpus and stelocyttarus calyptodomous nests—but he did not recognize that two species he placed in Parachartergus also have that condition, namely pusillus and ater (de Saussure).10 Ducke (1905a, 1905b) recognized four species groups in Parachartergus, only one of which had the fourth labial palpomere either rudimentary or lacking. That was his first group for species now placed in Parachartergus or Chartergellus Bequaert—species that, as far as is known, all have the labial palpi three-segmented, never with a rudimentary fourth segment. Ducke's (1905a, 1905b) three other species groups within Parachartergus were all stated to have the fourth segment of the labial palpus very distinct. That is correct for two of the groups, which are species now placed in Pseudopolybia de Saussure, but incorrect for his fourth group, which consisted solely of pusillus.
In 1914, Ducke transferred pusillus to Pseudopolybia, which was raised to generic rank by R. von Ihering (1913: 227; Pseudopolybia had been described as a division of Polybia by de Saussure, 1863). The species did not receive much further attention, although it was misidentified by Bequaert (1944), and following him Richards (in Richards and Richards, 1951), who used the name for Leipomeles spilogastra (Cameron) (see Carpenter, 1999: 9). Then Richards (1978: 202) studied a specimen, which he designated as lectotype, and transferred the species to his new genus, Marimbonda.
Ducke's (1914) definitive generic classification for neotropical Polistinae recognized 20 genera, a number that was not augmented until Richards (1978), who recognized 24. Of Richards' four additional genera, two were elevations of subgenera described by Bequaert (1938: Chartergellus, Nectarinella), while two were new taxa, Marimbonda and Occipitalia. The latter was subsequently synonymized with Clypearia by Carpenter et al. (1996), who showed the single species included in Occipitalia to be intermediate in the morphological and nest architectural characters defining Clypearia. Other work subsequent to Richards (Raw, 1985; Carpenter and Wenzel, 1990; Carpenter et al., 2000) has led to a number of 22 presently recognized neotropical polistine genera. Synonymy of Marimbonda with Leipomeles now reduces that number to 21.
SYNONYMY
Leipomeles and Marimbonda share numerous morphological features that are certainly derived: the third segment of the labial palpus bearing a stout, curved bristle apically, reduction in the palpal formula to five-segmented maxillary palpi and three-segmented labial palpi, mesepisternum without dorsal groove, and first metasomal segment conically petiolate. The two genera are not keyed out together in Richards' key (1978: 10), but that is partly erroneous. Thus, the pronotal fovea is not “weak or absent” in Leipomeles; when present, it is as “strong” as other genera. Also contrary to the key, the anterior pronotal carina (“prominence”) is in fact acute, and the metanotum in fact has a dorsal surface. The secondary spiracular entrance and propodeal valvula do not differ between Leipomeles and Marimbonda. This leaves the cuticle, which is less punctured and more shining in Leipomeles than Marimbonda, the form of the metasomal petiole which is more elongate in Leipomeles (cf. figs. 3 and 4), and the derived hindwing venation in Marimbonda (fig. 5). None of these features is as significant as the features shared by the two genera, and all are of less significance than the features by which other genera are differentiated (see key, below). Moreover, greater variation in all of these features is found within other genera: sharper differences in punctation occur in, for example, Polybia, Protopolybia, and Ropalidia; the form of the metasomal petiole is much more variable in, for example, Mischocyttarus and Polybia, and the difference between Leipomeles and Marimbonda is less than occurs within Polistes; and the hindwing venation is much more divergent within Parapolybia and Polybioides (van der Vecht, 1966; Carpenter, 1999), with CuA vein separating from M+CuA distad of the insertion of the cu-a crossvein, instead of basad of the insertion of cu-a.
The chief justification for recognition of Marimbonda as a genus in any case was the asterlocyttarus nest architecture. Only the nest collected with the type series of M. albogrisea is definitely known; Wenzel (1998: 16) stated to have seen another nest he believed to be Marimbonda, but the identity was not certain. According to Martin Cooper (in litt.), who collected the nest identified in Wenzel (1998: fig. 23D) as Marimbonda, it was found in Colombia, and was abandoned. Cooper's initial identification was as a Leipomeles nest, but Wenzel assigned it to Marimbonda because the comb was sessile. As will be seen, that is no longer grounds for generic assignment.
As discussed by Wenzel and Carpenter (1994), the nests of Marimbonda and Leipomeles share several derived architectural components, including application of blots of colored pulp to the envelope, envelope with ridges parallel to the lines of construction, and envelope with a central, impressed, longitudinal furrow. That last character is evidently variably present within both genera according to Wenzel (1998), who keyed out both genera together. Leipomeles always nests on the undersides of leaves, whereas the nest of Marimbonda collected by Richards was on the trunk of a sapling. The putative nest of Marimbonda was on a leaf (see Wenzel, 1998: 16), but it may have been Leipomeles. That aside, the absence of a petiole on the comb of Marimbonda was the only consistent difference in nest architecture between Leipomeles and Marimbonda. The number of petioles has long been known to be variable in Leipomeles, from one to several, and their orientation with respect to the substrate (leaf veins) may be vertical or oblique. Petioles have always been thought to be present, but I have now seen nests of Leipomeles dorsata (Fabricius) that lack them. First, a nest was sent to me by James H. Hunt, along with adult wasps, collected in Costa Rica: Prov. Heredia, La Selva Biological Station, 18 Jan. 1991 (J. H. Hunt), nest no. CR91–2. The single comb was attached to the underside of a leaf along the midrib not by a petiole, but rather by a low fibrous ridge that ran along the entire length of the comb. A hole in the ridge may possibly indicate fusion of originally separate structures, but if so these were ribbonlike rather than petiolate. Second, I have seen a nest at the Universidade de São Paulo in Ribeirão Preto, collected by Gabriel A. R. Melo in Brazil: Amazonas, Beruri, R. Purus, 16 Oct. 1991 (G. A. R. Melo), no. M89, along with adult wasps. The single comb was on the underside of a leaf of Astrocaryum, attached along its entire surface. Moreover, he collected another nest of this species in the same locality, identically attached along the entire surface.
Thus, astelocyttarus combs may occur in Leipomeles, and the main justification for separating Marimbonda collapses. Accordingly, I am synonymizing these genera. The synonymy follows, and then a new key to the genera of Polistinae of the New World.
Genus Leipomeles Möbius
Leipomeles Möbius, 1856, Nester Gesell. Wespen: 25, genus.
Type species: Leipomeles lamellaria Möbius, 1856 [= Polistes dorsata Fabricius, 1804], by monotypy.
Lipomeles Schulz, 1906, Spolia Hym.: 232. Un justified emendation of Leipomeles Möbius.
Marimbonda Richards, 1978, Social Wasps Am.: v, 10, 201, genus. new synonymy. Type species: Marimbonda albogrisea Richards, 1978, by original designation.
KEY TO THE GENERA OF NEW WORLD POLISTINAE
1. Metasomal segment I subsessile, funnel-shaped in dorsal view (fig. 7); propodeum with orifice dorsally acute (fig. 7); pronotum with a carina posterior to the lateral fovea (fig. 9)
— Metasomal segment I differently shaped, in dorsal view basally petiolate to long-petiolate or sessile with width greater than length (figs. 3–4, 8, 27, 29, 45–47); propodeum with orifice usually more broadly rounded dorsally (figs. 8, 45); pronotum if with carina and lateral fovea, then the carina is anterior to the fovea (fig. 10) or the two are not closely approximated (fig. 18)
2. Mid- and hindtarsi with third and fourth segments asymmetrical, inner lobe longer than outer lobe (fig. 11); forecoxa not dorsolaterally produced (fig. 13); metasomal segment I petiolate, not abruptly expanding posteriorly, in dorsal view with width half or less that of II, and at least twice as long as wide (fig. 8)
— Mid- and hindtarsi with third and fourth segments symmetrical (fig. 12); forecoxa usually dorsolaterally produced (fig. 14); metasomal seqment I variously shaped
3. Third labial palpomere with stout, recurved bristle apically (figs. 1, 2); female clypeus with square lateral lobes and sharply pointed apex (fig. 42)
— Third labial palpomere without bristle; female clypeus variable
4. Head with occipital carina present (figs. 15– 17)
— Head without occipital carina (fig. 18–19).
5. Pronotum with dorsal carina sinuous in lateral view (figs. 10, 17); forewing with prestigma about as long as pterostigma (fig. 20)
— Pronotum without sinuous carina (figs. 16, 18–19); forewing with prestigma shorter than pterostigma (fig. 21)
6. Metanotum vertical posteriorly (fig. 18); scutellum angular in profile (fig. 18)
— Metanotum rounded, not vertical posteriorly (fig. 19); scutellum rounded in profile (fig. 19)
7. Mesepisternum without dorsal groove (fig. 18); maxillary palpi five-segmented and labial palpi (fig. 1) three-segmented
— Mesepisternum with dorsal groove present, at least as anterior trace (fig. 19); maxillary palpi six-segmented and labial palpi (fig. 2) four-segmented
8. Pronotum without lateral fovea (figs. 22, 23)
— Pronotum with lateral fovea (figs. 18, 40), sometimes shallow
9. Scutellum angled in profile (fig. 22); metanotum not projecting beyond scutellum posteriorly (figs. 22, 24); head with occipital carina present dorsolaterally (fig. 22); pronotum without pretegular carina (fig. 24)
— Scutellum rounded in profile (fig. 23); projecting beyond scutellum posteriorly (figs. 23, 25); head with occipital carina present or absent; pronotum with pretegular carina (fig. 25)
10. Propodeum with dorsolateral carinae (fig. 26); metanotum with medial tubercle (fig. 26), sometimes weak; head with occipital carina present dorsolaterally (fig. 23)
— Propodeum without carinae (figs. 27–29); metanotum without tubercle (figs. 27–29); head without occipital carina (fig. 29)
11. Forecoxa not dorsolaterally produced (fig. 13)
— Forecoxa dorsolaterally produced (fig. 14)
12. Propodeum posteromedially concave (fig. 28); body with some pale maculations, not metallic; head in lateral view with tempora narrower than eye at ocular sinus (figs. 48–49); clypeus longer than wide.
— Propodeum posteromedially convex (fig. 29); body without pale maculations, cuticle partly to entirely bluish metallic or yellowish with some bluish highlights; head in lateral view with tempora as wide or wider than eye at ocular sinus (figs. 16, 50); clypeus at most as long as wide
13. Clypeus apically emarginate (fig. 30); cuticle black, bare, and shining
— Clypeus apically bluntly pointed (fig. 31); cuticle not shining
14. Metasomal segment I in lateral view more flattened posteriorly (fig. 32), in dorsal view with maximum width 3× basal width
— Metasomal segment I in lateral view convex posteriorly (fig. 33), in dorsal view with maximum width 2× basal width
15. Scutum with posterolateral lamella present adjoining tegula (fig. 34)
— Scutum with posterolateral lamella absent anteriorly, not adjoining tegula (fig. 35)
16. Ocelli enlarged, separated from eyes by less than an ocellar diameter (fig. 36); hindwing with jugal lobe reduced (fig. 38)
— Ocelli normal, separated from eyes by more than an ocellar diameter (fig. 37); hindwing with jugal lobe normal, not reduced (fig. 39)
17. Mesepisternum with dorsal groove (fig. 40); female clypeus with square lateral lobes and sharply pointed apex (fig. 42)
— Mesepisternum without dorsal groove (fig. 41); female clypeus with lateral lobes rounded (figs. 37, 43, 44) and apex bluntly pointed (fig. 37) or rounded (fig. 43) or truncate (fig. 44)
18. Metanotum produced posteromedially into elongate lobe overlapping the propodeum (fig. 45)
— Metanotum without posterior lobe
19. Metasomal segment I in dorsal view with short petiole, apically abruptly broadened, wider than half the width of segment II (fig. 46); head in lateral view with tempora much narrower than eye at ocular sinus (fig. 48); pronotum with anteromedial fovea
— Metasomal segment I in dorsal view differently shaped, narrowly petiolate to sessile (fig. 47); head in lateral view with tempora at least as wide as eye at ocular sinus (fig. 49), usually wider (fig. 50); pronotum without anteromedial fovea
20. Lateral ocelli almost as far apart as distance from each to eyes (fig. 51); eyes bristled (fig. 51); metasomal segment I not petiolate in dorsal view
— Lateral ocelli closer to each other than either is to eyes (fig. 37); eyes usually not bristled; metasomal segment I usually petiolate in dorsal view
Acknowledgments
I am very grateful to Fernando Noll and Sidnei Mateus for showing me an astelocyttarus nest of Leipomeles, and especially to Gabriel Melo for collecting it. I am also grateful to Jim Hunt for collecting and sending to me the nest from Costa Rica. I thank Martin Cooper, Kurt Pickett, and John Wenzel for encouragement and comments, and two anonymous reviewers of the manuscript for corrections. The illustrations for the key were provided by Molly Rightmyer and Valerie Giles.
REFERENCES
Figs. 1–6. Neotropical Polistinae. 1–2. Labial palpus. 1. Chartergellus nigerrimus Richards. 2. Pseudopolybia compressa (de Saussure). 3–4. Metasoma in dorsal view. 3. Marimbonda albogrisea Richards. 4. Leipomeles dorsata (Fabricius). 5–6. Hindwing. 5. Marimbonda albogrisea Richards. 6. Leipomeles dorsata (Fabricius). All scale bars are 1.0 mm
Figs. 7–12. Neotropical Polistinae. 7–8. Metasomal terga I and II in dorsal view. 7. Polistes consobrinus de Saussure. 8. Mischocyttarus rotundicollis (Cameron). 9–10. Head and mesosoma in anterolateral view. 9. Polistes consobrinus de Saussure. 10. Parachartergus fraternus (Gribodo). 11–12. Hindtarsus. 11. Mischocyttarus rotundicollis (Cameron). 12. Brachygastra augusti (de Saussure). All scale bars are 1.0 mm
Figs. 13–18. Neotropical Polistinae. 13–14. Head and prothorax in oblique frontal view, with procoxa set off in lateral view. 13. Mischocyttarus rotundicollis (Cameron). 14. Brachygastra augusti (de Saussure). 15–16. Agelaia multipicta (Haliday). 15. Head and mesosoma in oblique posterior view. 16. Lateral view. 17. Parachartergus fraternus (Gribodo), posterolateral view. 18. Nectarinella championi (Dover), head and mesosoma in lateral view. All scale bars are 1.0 mm
Figs. 19–24. Neotropical Polistinae. 19. Pseudopolybia vespiceps (de Saussure), head and mesosoma in oblique lateral view. 20–21. Forewing. 20. Parachartergus fraternus (Gribodo). 21. Angiopolybia obidensis (Ducke). 22–23. Head and mesosoma in lateral view. 22. Brachygastra augusti (de Saussure). 23. Chartergus artifex (Christ). 24. Brachygastra augusti (de Saussure), oblique posterolateral view, with pronotum and tegula set off. All scale bars are 1.0 mm
Figs. 25–30. Neotropical Polistinae. 25. Epipona guerini (de Saussure), oblique posterolateral view, with pronotum and tegula set off. 26–29. Oblique posterolateral view. 26. Chartergus artifex (Christ). 27. Asteloeca traili (Cameron). 28. Clypearia sulcata (de Saussure). 29. Synoeca cyanea (Fabricius). 30. Epipona guerini (de Saussure), head in frontal view. All scale bars are 1.0 mm
Figs. 31–36. Neotropical Polistinae. 31. Asteloeca traili (Cameron), head in frontal view. 32–33. lateral view. 32. Asteloeca traili (Cameron). 33. Metapolybia cingulata (Fabricius). 34–35. Scutum and tegula in dorsal view. 34. Apoica pallens (Fabricius). 35. Polybia bistriata (Fabricius). 36. Apoica pallens (Fabricius), head in frontal view. All scale bars are 1.0 mm
Figs. 37–42. Neotropical Polistinae. 37. Polybia bistriata (Fabricius), head in frontal view. 38–39. Hindwing. 38. Apoica pallens (Fabricius). 39. Agelaia multipicta (Haliday), hindwing. 40–41. lateral view. 40. Angiopolybia obidensis (Ducke). 41. Protopolybia sedula (de Saussure). 42. Angiopolybia obidensis (Ducke), head in frontal view. All scale bars are 1.0 mm
Figs. 43–48. Neotropical Polistinae. 43–44. Head in frontal view. 43. Protopolybia sedula (de Saussure). 44. Charterginus fulvus Fox. 45–46. Dorsal view. 45. Protopolybia sedula (de Saussure). 46. Charterginus fulvus Fox. 47. Polybia depressa (Ducke), metasoma in dorsal view. 48. Charterginus fulvus Fox, head and mesosoma in lateral view. All scale bars are 1.0 mm
[1] Junior homonym of Cyclostoma Lamarck, 1799; replaced by Gyrostoma Kirby, 1828; now a subgenus of Polistes (Carpenter, 1996).
[2] Suppressed by Opinion 1051 of the International Commission on Zoological Nomenclature (1976); the name Angiopolybia Araujo, 1946, is now used for this genus.
[3] Synonymized with Ropalidia Guérin-Méneville, 1831, by Kojima (1997).
[4] Now a subgenus of Polybia (Richards, 1978).
[5] Junior homonym of Raphigaster Laporte, 1832; replaced by Belonogaster de Saussure, 1853.
[6] Synonymized with Ropalidia Guérin-Méneville, 1831, by Kojima (1997).
[7] Junior homonym of Pseudopolybia de Saussure, 1863; junior synonym of Kappa de Saussure, 1854, a subgenus of Mischocyttarus.
[8] Now a subgenus of Mischocyttarus (Richards, 1941).
[9] Preoccupied; the name has been replaced by Chartergellus amazonicus Richards, 1978.
