James M. Carpenter.
The phylogenetic system of the Gayellini (Hymenoptera: Vespidae: Masarinae).
Psyche 95:211-242, 1988.

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THE PHYLOGENETIC SYSTEM OF THE GAYELLINI (HYMENOPTERA: VESPIDAE; MASARINAE)*
Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138
The Gayellini is one of the two tribes of Masarinae (Carpenter, 1981). Endemic to the Neotropics, the majority of the species are Patagonian, but one ranges as far north as Mexico. With ten de- scribed species, the group is far less speciose than its sister-tribe Masarini, which has over 200 described species (cf. Richards, 1962), and most species are rarely collected. These wasps have a very dis- tinctive appearance among Vespidae (Fig. l), and their taxonomic history has been more turbulent than any other higher vespid taxon. Although the phylogenetic placement of the group as a whole has now evidently been settled (Carpenter, 1981), no study has been made of the species. The current generic classification is fragmented, and there have been no keys to all of the taxa. In this paper, I investigate the phylogenetic relationships of the species, and present a revised generic classification along with keys to all taxa. Saussure (1852-58) placed Gayella in the Section "Anomalop- tkres" of the "Eumkniens" because the forewing recurrent veins (m-cut-2) are received in separate cells (Fig. 6), as in the other genera placed in this section (Raphiglossa, and Stenoglossa = Psili- glossa). In other vespids he studied both veins were received by the second submarginal cell. Ashmead (1902a) described the subfamily Raphiglossinae (in his Eumenidae) for this group, but by that time other genera had been described which had the diagnostic character of the recurrent veins. These were Euparagia and Paramasaris, both considered probable masarines by their authors (Cresson, 1879, and Cameron, 1901, respectively). Ashmead (1902b) proposed the tribe Euparagiini in his Masaridae for these two genera. So the recurrent *Manuscript received by the editor September 28, 1988 21 1




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212 Psyche [VOI. 95
veins were no longer uniquely diagnostic. Bequaert (1 9 18) ques- tioned whether Gayella belonged in the Raphiglossinae, since the longitudinal plaiting of the forewings "is very obsolete" in the genus, and Bradley (1922) placed it in its own subfamily in Vespidae s. I. Bequaert (1928) transferred Paramasaris to this subfamily, based on its possession of the characteristic hindwing venation of Gayella (Fig. 3). Richards (1962) included the Gayellinae in his Masaridae, but his dendrogram showed Euparagiinae as more closely related to the subfamily Masarinae. I (Carpenter, 1981) demonstrated that Richards' Masaridae was a paraphyletic group, since Euparagiinae is the sister-group of Vespidae as a whole. Four synapomorphies were adduced which showed a sister-group relationship between gayellines and masarines: presence of hypostomal apodemes, loss of the midfemur basal ring, loss of the scutal lamella and provisioning with nectar and pollen. Gayellines and masarines were treated as tribes in an expanded Masarinae, the system followed here. Gayella was originally described as monotypic for (7. eumenoides by Spinola (1851). Saussure (1855 in Vol. 3 of Etudes), Willink (1956, 1963) and Willink and Ajmat de Toledo (1979) added five species. The latter paper provided a key to the six species, however I believe that the key given here will be easier to use. Paramasaris was also originally described as monotypic, for P. fuscipennis Cameron. Cameron (1904) later described a new genus Zethoides (non Zethoides Fox, 1899; Plesiozethus Cameron, 1905, and Metaze- thoides Schulz, 1906, are replacement names) for 2. flavolineatus, which differed from Paramasaris in having only two (Fig. 5), as opposed to three (Fig. 6), submarginal cells. Zavattari (1912) ques- tioned whether Cameron had described this character correctly, and Bradley (1922) suspected that Plesiozethus was a synonym of Para- masaris. This was confirmed by Bequaert (1928), who showed that the number of submarginal cells was variable, and synonymized flavolineatus with fuscipennis. Giordani Soika (1974) described two new species in the genus, but provided no key. He also described a new genus, Paragayella, monotypic for the new species Paragayella richardsi. I consider this genus a synonym of Paramasaris, as dis- cussed below. I also give the first key to species of the group. All of the species have been examined. Types of Paramasaris have been seen; treatment of Gayella follows Willink's concepts. Com-



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Carpenter - Gayellini
Fig. 1 .
Gayella eumenoides, 9.




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214 Psyche [vo~. 95
plete label data for all material of Paramasaris are listed under taxonomic notes for each species; for the relatively better known Gayella only provinces are noted. Acronyms for collections are listed below, along with the name of the individuals who provided the material where this was borrowed.
AMNH
BMNH
CAS
CP
IML
IPC
MCZ
MF
MNHN
UCD
USNM
American Museum of Natural History, New York (M. S. Favreau)
British Museum of Natural History, London (M. C. Day, C. R. Vardy)
California Academy of Sciences, San Francisco (W. J. Pulawski)
Charles Porter personal collection
Instituto Miguel Lillo, Tucuman (A. Willink) Instituto Pedagbgico de Chile
Museum of Comparative Zoology, Cambridge M. A. Fritz personal collection
Museum National d'Histoire Naturelle, Paris University of California, Davis (P. S. Ward) U.S. National Museum of Natural History, Washington (A. S. Menke)
Morphological terminology follows Carpenter (1981), except that I have adopted Snelling's (1986) more descriptive terms "preoccipi- tal" and "po~tocular'~ for the carinae previously termed "dorsal occipital keel" and "ventral occipital keel" (Richards, 1962). Detailed examination of the labiomaxillary complex and male geni- talia was made by dissection of these structures, clearing slightly in cold lactophenol, and examination in glycerin. Measurements were made with an ocular micrometer. Illustrations were made with a Wild M-400 photomacroscope employing Kadak TMAX 400 film. Cladistic analysis (Hennig, 1966) was performed for all the features discussed in this paper. Outgroup taxa include Masarini and Euparagiinae, with reference to other Vespidae also occasionally made.
Subfamily characters
First I discuss some morphological features important in higher- level vespid relationships, before turning to consideration of the phylogenetic relationships among the species. Autapomorphies of



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Carpenter - Gayellini 215
G. mutilloides
G. luispenai
G. patagonica
G. reedi
G. araucana
P. brasiliensis
P. cupreus
P. f uscipennis
Fig. 2.
Cladogram of the species of Gayellini.
\ G. eumenoides
P. richardsi




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216 Psyche [vol. 95
the Gayellini listed by Carpenter (198 1) include the hindwing with CUI diverging from M+Cul far distad of the insertion of cu-a (Fig. 3), the clypeus with the dorsal margin bisinuate (Fig. 12), the first metasomal tergum and sternum fused and metasomal segments after I1 retractile (the latter two convergent with other vespids). Some other autapomorphies are mentioned below. Forewing discal cell. Carpenter (198 1 : 14) noted that the discal cell is shorter than the submedian in Paramasaris. This is also the case in Gayella (Fig. l), and this should be considered a reversion from the state of an elongate discal cell in other Vespidae (Fig. 4; cf. Carpenter, 1981), and thus an autapomorphy of Gayellini. Forewing radial region.
The variation in the number of sub-
marginal cells in Paramasaris was alluded to above. Besides fusci- pennis, I have seen loss of r-m2 producing two submarginal cells in several specimens of brasiliensis (including the allotype and para- type, Fig. 5). The placement of m-cu~ varies as well, sometimes meeting M at the fork where RS diverges. But this is not correlated with number of cells, and most specimens have the usual condition of RS diverging first (Figs. 5-6). In addition, the specimen of Para- gayella richardsi from Formoso, Brazil, has a very small adventi- tious cell at the junction of r-m3 and RS on one wing (Fig. 6), and both Goias specimens have an adventitious vein spur arising from the middle of r-m3 (Fig. 6).
Clypeus. The clypeus is narrower than its height in all species, particularly in males (Figs. 1 1 - 17). Richards (1 962:46) stated that the reverse is true in Paramasaris, perhaps a lapsus. This is not the usual state in Vespidae, and is perhaps apomorphic, although the degree of narrowing varies in the tribe. Occipital carinae. Gayellini have both the postocular and preoccipital carinae in the groundplan, contrary to Richards (1962:12). The postocular carina is reduced in length, and may be present only as a trace just ventral to the eye in Gayella, but is typically obvious in the eumenoides group. The carinae are almost confluent in many specimens of eumenoides and araucana, sepa- rated by only a slight gap (Fig. 18). The "complete" carina in Para- masaris (Richards, 1962:46; Fig. 19) is evidently produced by confluence of the postocular with the preoccipital carina, as occurs in some Masarini (Snelling, 1986) and probably other Vespidae (Carpenter, 1988). The postocular carina is effaced in Paragayella (Fig. 20).




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Carpenter - Gay ellini 217
Figs. 3- 10. 3. Guyella araucana, 7k. Hindwing. 4. Paragia decIpMas alicke, 4X. Wings. 5-6. Wings, 6X.
5. Paramaswis brmifiensis.
6. P. richardsi. Submar-
ginal cells are numbered.
7- 10. Lateral view of pronoturn. 7. G. araucana, IPX. 8. Eupatagia scutellaris, !OX.
9. Jschnocoelia robusta, 6X.
10. Yespa affinis,
5X. ac: adventitious cell; ap: anterior pronotal canna; av: adventitious vein; d: discal cell;/' pronotal fovea; p: pronotal carina; rv: recurrent veins,



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218 Psyche [vo~. 95
Acroglossal buttons.
As noted by Carpenter (1981), Richards
(1962) incorrectly stated that acroglossal buttons are lacking from the ligula of Gayellini. The only species in which they are entirely lacking are Pararnasaris fuscipennis and cupreus (Fig. 21). These structures are also absent from the glossa of Paramasaris brasilien- sis, but are present on the paraglossae. This is a transformation series in reduction. Other gayellines have the buttons on both the glossa and paraglossae (Fig. 22), including Paragayella richardsi, the sister-group of Paramasaris, and P. brasiliensis is the sister- group of fuscipennis + cupreus (Fig. 2). Hypostomal apodemes. These are present in all species, which supports the interpretation of synapomorphy with Masarini. They are always very narrow (Fig. 23).
Pronotal carina. Paramasaris and Paragayella are notable for having two parallel carinae on the pronotum. One is present at the anteroventral margin of the pronotum; the other is posterior to this and runs towards the humeri and dorsum of the pronotum (Figs. 29-31, 40). The second carina shows a transformation series in development, ranging from short lateral sections only (Paragayella, Figs. 24, 29), to extending to the dorsum (Paramasaris brasiliensis, Figs. 25, 30), to complete across the dorsum (P. cupreus and fuscipennis, Fig. 26). This series apparently corresponds to the phylogenetic relationships among these species (Fig. 2). Gayella has only the anterior carina (Fig. 7). Euparagiinae also has only the anterior carina (Fig. 8), although the humeri are somewhat raised in scutellaris. In Masarini the anterior carina is usually blunt, and a lateral carina on the humeri may be present (Fig. 37). In all these groups, the anterior carina precedes a groove which is frequently crenate (secondarily reduced in various Masarini). The situation is different in other Vespidae. In Polistinae, the structure termed the "pronotal prominence" (Richards, 1978) is probably homologous with the anterior carina. Although often blunt, it is frequently carinate, and lies at the anteroventral margin of the pronotum (Fig. 38). It precedes the pronotal fovea, which is sometimes set in a deep depression; there is no lateral groove. In the groundplan, there is also a carina on the dorsum (Carpenter, 1989). This second carina is usually quite short laterally, and may closely approach the anterior carina (Fig. 38). In Polistes the second carina extends almost to the ventral pronotal margin in many species, and the fovea, which is anterior to this carina, is not preceded by a



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Carpenter - Gayellini
Figs. 1 1 - 1 9. I I- 1 2. Clypeus, 9. f 1. Paramusaris richurdst, 19X. I2 P. brasi- liensis, 27X. 13-16. Frontalviewofhead,Q. 13, P.cup~eus. 14X. 14. Gayella mutihides, 5X. 15. G, eumenoides, 7X 16. G. uraucanar 10X. 17. ff. reedi 8, )OX. Frontal view of head.
18-19. Lateral view of head. 18. G. waucaw 15X. 19. P. brasiliemis, 21X. cc: apical clypeal carinae; g: gap between mandibular teeth; po: postocular carina; pr: preoccipital carina.



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220 Psyche [vo~. 95
"prominence" (Fig. 39). Richards (1973) confused the dorsal carina in Polistes, behind the fovea, with the anterior carina in other polistines, in front of the fovea. Eumeninae, which also have a fovea, also have a carina in front of the fovea (Fig. 9), which continues across the dorsum in the groundplan (Carpenter and Cumming, 1985). The single carina may be a composite structure, derived from a state resembling certain polistines with two closely approximated carinae (Fig. 38). This is also the case in the ground- plan of Vespinae (Carpenter, 1987), where there is a single carina, preceding the pronotal fovea and running across the dorsum (Fig. 10). Stenogastrinae have a highly modified pronotum and lack a posterior carina and fovea (Carpenter, 1988), but have a blunt ridge anteriorly that may correspond to the anterior carina (Fig. 32). Thus, an anterior carina is clearly an ancestral vespid character, but considering its diverse form, the posterior carina may have evolved multiple times. The alternative interpretation, that it evolved once (in the ancestor of all vespids except Euparagiinae), requires secondary losses within Gayellini (Paragayella, Paramasa- ris and Gayella independently) and Stenogastrinae. The interpreta- tion of nonhomology is more parsimonious, and accords with the apparent transformation series in Paramasaris. In any case, the separate posterior carina has also been lost on numerous occasions within the Masarini and Polistinae (Carpenter, 1989, and in prep.), and the possibly composite carina has also been lost several times within Eumeninae and Vespinae (Carpenter, 1987; Carpenter and Cumming, 1985). Secondary loss also applies to the pronotal fovea, present in the groundplan of Eumeninae, Polistinae and Vespinae. It has been lost multiple times within Polistinae (Richards, 1978; Carpenter, 1989), and probably also in Stenogastrinae. Presently available morphological and behavioral evidence supports a sister- group relationship between Stenogastrinae and Polistinae + Vespi- Figs. 20-28.
20. Paramasaris richardsi, 17X. Lateral view of head. 21 -22. Ven-
tral view of ligula. 21. P. fuscipennis, 25X. 22. Gayella luispenai, 16X. 23. G. mutilloides, 14X. Ventral view of head, mouthparts removed. 24-28. Dorsofrontal
view of pronotum. 24. P. richardsi, 9X. 25. P. brasiliensis, 11~. 26. P. cupreus, 13X. 27. G. eumenoides 8, 10X. 28. G. araucana Q, 8X. ab: acroglossal buttons; ap' anterior pronotal carina; gl: glossa; h: hypostomal apodeme; hp: humeral projec- tion; pg: paraglossa; pip: posterior lingual plate; pp: posterior pronotal carina; pr: preoccipital carina.




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198231 Carpenter - Gayelhi 22 1




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222 Psyche [vo~. 95
nae (Carpenter, 198 1 ? 1988, 1989)? thus requiring an inference of loss in Stenogastrinae.
Hind coxal carina.
Richards (1962) made contradictory state- ments concerning the presence of this feature in Gayellini (cf. p. 15 and 44). This was initially followed by Carpenter (1981), but cor- rected by Carpenter and Cumming (19851907). All Gayellini lack this carina, a primitive condition.
Claws. Richards (1962:44) erroneously characterized the claws of Gayella as simple, and Carpenter (198 1:26) initially followed this (corrected in Carpenter and Gumming? 1985:907). In fact, the claws are toothed in all species of Gayellini (variable in G. rnutilloides). This is the plesiomorphic condition in Vespidae. Male genitalia. I have examined the genitalia of all species except Paramasaris cupreus and Paragayella richardsi) where the males are still unknown. In the groundplan of the tribe, the aedea- gus is broadly rounded apically9 the digitus is a prominent triangu- lar lobe that is desclerotized ventrally, the cuspis is a small lobe completely fused to the lamina volsellaris, and the parameral spines are long and sharply pointed (Figs. 56-63). Figure 39 of Richards (1962), showing a large? triangular cuspis and rounded digitus in Gayella araucana, is incorrectly labelled. What is there termed cus- pis is actually the digitus? and the structure labelled as digitus must be the aedeagus (cJ Fig. 58). This figure was the reason I previously was unable to characterize the groundplan of the volsella in the tribe (Carpenter, 1981:26), as I had not seen that species at the time. Within genera, the genitalia are relatively uniform? with species dif- fering in details (especially of the volsella); however, there are some consistent differences between the genera. These are discussed below.
Pararnasaris
Giordani Soika (1974) characterized Paragayella as related to Gayella, and stated (my translation): "This genus appears at first sight a Gayella by the general aspect and dimensions." The type material I have seen he even labelled as "Gayella richardsi. "In fact, Paragayella is not really even superficially similar to Gayella. Para- gayella lacks some of the apomorphies shared by the species of Pararnasaris, and for some other derived traits which Paramasaris and Paragayella share the latter has a less developed state. Thus it shares some primitive similarity with Gayellay which of course indi-



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19881 Carpenter - Gayellini 223
cates nothing about phylogenetic relationship (Hennig, 1966). On the other hand, Paragayella shares several clear synapomorphies with Paramasaris. These include the forewing with r-m3 more or less straight (Figs. 5-6; sinuous in Gayella and other Vespidae, Figs. 1,4)? the pronotum with two carinae (Figs. 29-3 1,40; one in Gayella Fig. 71, the metanotum with a longitudinal median carina (Figs. 33, 46; none in other Masarinae)? and the metasoma petiolate (tergum I in dorsal view at least twice as long as wide and half the width of tergum II? Figs. 35? 40-41; it is differently shaped in Euparagiinae, Masarini and Gayella, Figs. 1, 43-45). Paragayella is the sister- group of Paramasaris. Autapomorphies of Paragayella include the reduced postocular carina (Fig. 20) and the transversely carinate metanotum (Fig. 46).
The three species of Paramasaris share numerous synapomor- phies. The postocular and preoccipital carinae are apparently con- fluent (Fig. 19). These carinae are separated in other Gayellini, and the postocular carina reduced in several species (Paragayella, the Gayella mutilloides group). The mandibles are tridentate with the proximal teeth separated from the apical one by a gap (Fig. 13). The mandibles are quadridentate in females of Paragayella and Gayella (Figs. 11, 14-15), and tridentate in males of the latter (Fig. 17; Richards, 1962:44, erroneously characterized the mandibles of Paramasaris as quadridentate and those of Gayella as simple); there is no gap. Quadridentate mandibles is the groundplan state of most of the Vespidae (Carpenter, 1981), although Euparagiinae has bidentate mandibles. The glossa is shortened and lacks acroglossal buttons, the paraglossae are broadened? and the posterior lingual plate is cordate in shape (Fig. 21). The posterior lingual plate is slightly broadened in other gayellines? but the length of the structure still exceeds its width (Fig. 22). The clypeus is broadly truncate (Fig. 121, which is here treated as derived, convergent with the ground- plan of Masarini. Paragayella has the clypeus narrowly truncate (Fig. 1 l)? as in Euparagiinae, which is considered the primitive state, A broad truncation seems most simply interpreted as derived from a narrow emargination, as does the pointed clypeus of Gayella (Figs. 14-17). The posterior carina of the pronotum extends further dor- sad in Pararnasaris (Figs. 25-26) than Paragayella (Fig. 24), a further apomorphic development. The propodeum has oblique carinae more or less developed (Figs. 31? 331, a unique trait in



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224 Psyche [VOI. 95
Figs, 29-36.
29-32. Lateral view nf pronoturn and mesepisternum. 29. Pam+ masaris richurdsi, 13Y. 30. P. hrmiliensb, f4X. 31. P. fuscipennis, 17X. 32. Pnrischnogmter meI[vi, l6X. 33-34, Oblique lateral view of propodeurn. 33. P. fuscipennis. 17X. 34. P. cupreus. 1SX. 35. P, brmiliensis holotype, 9X.
Lateral view of metasoma.
36. Ga,sella reedi 8, 3X. Lateral view. op: anterior pronotat carina; at: anterior t runcation of metasomal tergum I; cg: carina delimiting



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19881 Carpenter - Gayellini 225
Masarinae. The first metasomal tergum has a blunt posterior ridge that is continued anterolaterally and drawn out into projections posterolaterally (Figs. 35, 401, a feature unique in Vespidae. The tergum is also strongly truncate anteriorly (Fig. 35). The second tergum has a median longitudinal ridge (Figs. 40, 42), which how- ever is variably developed in brasiliensis (strong in the male and not developed in the female, Fig. 35). A longitudinal ridge is found elsewhere in Vespidae only within Eumeninae (Cyphomenes, where it is anterior). Finally, the parameral spines of the male genitalia are extremely elongate in brasiliensis and fusc@ennis, being longer than the parameres and extending far beyond the apex of the aedeagus (Fig. 56). This is apparently a derived condition; in Gayella, Eupa- ragiinae and Masarini the spines are shorter than the parameres and extend little beyond the aedeagus (Figs. 57-63). Males of cupreus are predicted to share this synapomorphy, and possibly also Paragayella.
Within Paramasaris, cupreus and fuscipennis are sister-groups. This is shown by the paraglossae also lacking acroglossal buttons (Fig. 211, the female clypeus with a pair of short apical carinae (Fig. 131, the second carina of the pronoturn more complete dorsally (Fig. 261, and the longitudinal carina on tergum I1 well developed in females (Figs. 40, 42). Autapomorphies of the species are: for cupreus the propodeal median groove delimited by more lamellate carinae (Fig. 34); for fuscipennis the oblique propodeal carina better defined (Fig. 331, and the dorsal groove and scrobal furrow of the mesepisternum broader and deeper (Fig. 31, cJ with 29-30, 40). I have not discovered any autapomorphies of brasiliensis. Since Paragayella is the sister-group of Paramasaris, recognition