ABSTRACT
Advances in potter wasp systematics have been achieved recently, with classificatory changes resulting from analyses based upon large scale molecular datasets. For the Neotropics, recent hypotheses point to the occurrence of an exclusive clade recognized within the tribe Eumenini. In this group, several contributions regarding taxonomy and systematics have been proposed in the last five years, including the genus Alphamenes. This taxon contains seven described species whose distribution is exclusively Neotropical. Females are morphologically homogeneous, and characters related to copulatory organs are useful in male diagnosis. This contribution forms the first phylogenetic approach to include all species of Alphamenes, hence the first to strongly test for group monophyly. Our cladistic results recovered Alphamenes as a monophyletic group supported by male genital features. Relationships among included species also rely upon genitalic characters, highlighting the importance of these attributes for eumenine systematics. Recent phylogenetic investigations applied to the Neotropical fauna of potter wasps represent desirable advancements towards a natural classification for the group.
Keywords:
Male genitalia; Minixi; Systematics; Taxonomy
Introduction
Phylogenetic studies based upon molecular data have recently changed our understanding of the evolutionary relationships within the eumenines, the so-called potter wasps (Bank et al., 2017Bank, S., Sann, M., Mayer, C., Meusemann, K., Donath, A., Podsiadlowski, L., Kozlov, A., Petersen, M., Krogmann, L., Meier, R., Rosa, P., Schmitt, T., Wurdack, M., Liu, S., Zhou, X., Misof, B., Peters, R., Niehuis, O., 2017. Transcriptome and target DNA enrichment sequence data provide new insights into the phylogeny of vespid wasps (Hymenoptera: Aculeata: Vespidae). Mol. Phylogenet. Evol. 116, 213-226.; Hines et al., 2007Hines, H.M., Hunt, J.H., O'Connor, T.K., Gillespie, J.J., Cameron, S.A., 2007. Multigene phylogeny reveals eusociality evolved twice in vespid wasps. Proc. Natl. Acad. Sci. U.S.A. 104, 3295-3299.; Piekarski et al., 2018Piekarski, P.K., Carpenter, J.M., Lemmon, A.R., Lemmon, E.M., Sharanowski, B.J., 2018. Phylogenomic evidence overtunrs current conceptions of social evolution in wasps (Vespidae). Mol. Biol. Evol. 35, 2097-2109.). These studies contradict our traditional view of the vespid wasp – and by extent, relationships within the eumenines – phylogenetic relationships based on morphological data (Carpenter, 1982Carpenter, J.M., 1982. The phylogenetic relationships and natural classification of the Vespoidea (Hymenoptera). Syst. Entomol. 7, 11-38.; Hermes et al., 2014Hermes, M.G., Melo, G.A.R., Carpenter, J.M., 2014. The higher level phylogenetic relationships of the Eumeninae (Insecta, Hymenoptera Vespidae), with emphasis on Eumenes sensu lato. Cladistics 30, 453-484.) and a total evidence approach (Pickett and Carpenter, 2010Pickett, K.M., Carpenter, J.M., 2010. Simultaneous analysis and the origin of eusociality in the Vespidae (Insecta: Hymenoptera). Arthropod Syst. Phylo. 68, 3-33.), rendering Eumeninae paraphyletic.
In the Neotropical region, advances in the systematics of the group were achieved only recently (e.g. Hermes and Carpenter, 2012Hermes, M.G., Carpenter, J.M., 2012. Are Monobia and Montezumia monophyletic? A cladistic analysis of their species groups based on morphological data (Hymenoptera, Vespidae, Eumeninae). Am. Mus. Nov. 3733, 1-23.; Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.). Despite the fact that the tribal classification proposed by Hermes et al. (2014)Hermes, M.G., Melo, G.A.R., Carpenter, J.M., 2014. The higher level phylogenetic relationships of the Eumeninae (Insecta, Hymenoptera Vespidae), with emphasis on Eumenes sensu lato. Cladistics 30, 453-484. was challenged by molecular studies, their results indicate that a subgroup of Eumeninae may indeed be an exclusive Neotropical clade. This clade was recognized as "Clade 4" in their contribution, which contains the majority of the taxonomic diversity within their tribe Eumenini.
Within the Neotropical Eumenini sensu Hermes et al. (2014)Hermes, M.G., Melo, G.A.R., Carpenter, J.M., 2014. The higher level phylogenetic relationships of the Eumeninae (Insecta, Hymenoptera Vespidae), with emphasis on Eumenes sensu lato. Cladistics 30, 453-484., some contributions encompassing a wide range of taxa were recently published (e.g. Grandinete et al., 2015Grandinete, Y.C., Hermes, M.G., Noll, F.B., 2015. Systematics and phylogeny of the Neotropical Pachymenes de Saussure and Santamenes Giordani Soika (Hymenoptera, Vespidae Eumeninae). Syst. Entomol. 40, 365-384.; Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.). The eumenine classification within this region has been continually investigated under cladistic approaches, with generic synonymy resulting in the cases of Pachymenes de Saussure, 1852 and Santamenes Giordani Soika, 1990 (Grandinete et al., 2015Grandinete, Y.C., Hermes, M.G., Noll, F.B., 2015. Systematics and phylogeny of the Neotropical Pachymenes de Saussure and Santamenes Giordani Soika (Hymenoptera, Vespidae Eumeninae). Syst. Entomol. 40, 365-384.) and MinixiGiordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420. and PachyminixiGiordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420. (Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.). Regarding the latter, a special case of a closely related taxon, namely Alphamenes van der Vecht, 1977, remains to be carefully addressed. Sexual association for members of this genus is particularly difficult, with females of some species being morphologically similar (Giordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420.; Oliveira et al., 2017Oliveira, L.A., Ferreira, W.D., Inez, T.G., Hermes, M.G., 2017. Contributions to the taxonomy of Minixi Giordani Soika and Alphamenes van der Vecht (Hymenoptera, Vespidae Eumeninae), with an identification key to the species of the latter. Zootaxa 4317, 530-540.). Also, no phylogenetic study to date have included all species of Alphamenes, thus its monophyly is still open to further investigation.
The present contribution enhances our knowledge about the Neotropical fauna of eumenine wasps, aiming at testing the monophyly of Alphamenes and recovering the phylogenetic relationships among its species. This forms our final contribution to a series of papers dealing with the systematics of Alphamenes and Minixi (Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.; Oliveira et al., 2017Oliveira, L.A., Ferreira, W.D., Inez, T.G., Hermes, M.G., 2017. Contributions to the taxonomy of Minixi Giordani Soika and Alphamenes van der Vecht (Hymenoptera, Vespidae Eumeninae), with an identification key to the species of the latter. Zootaxa 4317, 530-540.), whose generic status have been traditionally questioned by eumenine workers in the last couple of decades (see Carpenter and Garcete-Barrett, 2003Carpenter, J.M., Garcete-Barrett, B.R., 2003. A key to the Neotropical genera of Eumeninae (Hymenoptera: Vespidae). Bol. Mus. Nac. Hist. Nat. Paraguay 14, 52-73.).
Material and methods
Material examination and morphological studies
We examined 376 specimens belonging to Alphamenes and Minixi (Table 3 in Supplementary Material). The specimens were borrowed from the following institutions: AMNH – American Museum of Natural History, New York, USA (Dr. James M. Carpenter); MCZ – Museum of Comparative Zoology, Harvard University, Cambridge, USA (Dr. Jignasha Rana); MNHN – Muséum National d'Histoire Naturelle, Paris, France (Dr. Claire Villemant); MNHNPY – Museo Nacional de Historia Natural del Paraguay, San Lorenzo, Paraguay (Dr. Bolívar R. Garcete-Barrett); CMNH – Carnegie Museum of Natural History, Pittsburgh, USA (Dr. John Rawlins); NHM – Natural History Museum, London, England (Dr. Gavin Broad); ZMHB – Museum für Naturkunde der Humboldt-Universität, Berlin, Germany (Dr. Frank Koch); MSNVE – Museo di Storia Naturale di Venezia, Venice, Italy (Dr. Marco Uliana); HYMSJRP – Coleção de Hymenoptera, Departamento de Zoologia e Botânica, Instituto de Biociências, Letras e Ciências Exatas, Universidade Estadual Paulista ‘Júlio de Mesquita Filho', São José do Rio Preto, Brazil (Dr. Fernando B. Noll); UFMG – Coleções Taxonômicas da Universidade Federal de Minas Gerais, Belo Horizonte, Brazil (Dr. Fernando A. Silveira); DZUP – Coleção de Entomologia Pe. Jesus Santiago Moure, Departamento de Zoologia da Universidade Federal do Paraná, Curitiba, Brazil (Dr. Gabriel Melo); CEUFLA – Coleção Entomológica da Universidade Federal de Lavras, Lavras, Brazil (Dr. Marcel G. Hermes).
The morphological study of the specimens was carried out with the stereoscopic microscope Leica S8 APO. Images of the structures of phylogenetic and taxonomic interest were obtained with a Leica 104 45929 digital câmera coupled to the Leica DFC295 stereoscope microscope. Terminology for male genitalia followed Bitsch (2012)Bitsch, J., 2012. Morphologie comparée des derniers segments du gastre et des genitalia mâles des Vespidae. 1. Sous-famille des Eumeninae (Hymenoptera). Bull. Soc. Entomol. France 117, 199-218. and for external morphology Carpenter and Garcete-Barrett (2003)Carpenter, J.M., Garcete-Barrett, B.R., 2003. A key to the Neotropical genera of Eumeninae (Hymenoptera: Vespidae). Bol. Mus. Nac. Hist. Nat. Paraguay 14, 52-73.. The male genitalia was detached from the metasoma and clarified on 10% NaOH solution for 24–36 h, then neutralized with acetic acid, washed in water and stored in glycerol.
Cladistic analyses
For the reconstruction of the phylogenetic relationships, all recognized species within Alphamenes and Minixi were included as ingroup (sensuHermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.). In the case of M. atrumSelis, 2017Selis, M., 2017. A new species of Minixi Giordani Soika, 1978 from Argentina (Hymenoptera Vespidae, Eumeninae). Linzer Biol. Beitr. 49, 687-690., the information was extracted from the literature (Selis, 2017Selis, M., 2017. A new species of Minixi Giordani Soika, 1978 from Argentina (Hymenoptera Vespidae, Eumeninae). Linzer Biol. Beitr. 49, 687-690.). We based our analyses upon our previous matrix (Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.), with the inclusion of all species of Alphamenes and adjusting character coding accordingly. This resulted in a matrix with 42 morphological characters, an increase of eight characters from Hermes and Oliveira (2016)Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.. Species of the closely related LaevimenesGiordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420. and Pachymenes de Saussure, 1852 were included in the analyzes. The inclusion of these taxa in the analyzes aimed at testing the monophyly of the ingroup genera (mainly Alphamenes and Minixi). Despite the fact that Minixi was already investigated phylogenetically (Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.), we opted for including its species to perform a stronger test (more ingroup taxa and more characters) of the monophyly of the taxa under scrutiny. For the rooting of the trees the outgroup method was used (Nixon and Carpenter, 1993Nixon, K.C., Carpenter, J.M., 1993. On outgroups. Cladistics 9, 413-426.) and the species Monobia angulosa de Saussure, 1852 was selected.
Fitch's parsimony (Fitch, 1971Fitch, W.N., 1971. Toward defining the course of evolution: minimum change for a specified tree topology. Syst. Zool. 20, 406-416.) was the optimization criterion used in the present study. Character matrix construction was carried out with Winclada v. 1.00.08 (Nixon, 1999–2002Nixon, K.C., 1999–2002. Winclada, ver. 1.0000. Published by the author. Ithaca, New York.). Heuristic searches for the most parsimonious cladograms were performed with TNT v. 1.5 (Goloboff and Catalano, 2016Goloboff, P.A., Catalano, S.A., 2016. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32, 221-238.), using implied character weighing (Goloboff, 1993Goloboff, P.A., 1993. Estimating character weights during tree search. Cladistics 9, 83-91.; Goloboff et al., 2008Goloboff, P.A., Carpenter, J.M., Arias, J.S., Esquivel, D.R.M., 2008. Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics 24, 758-773.). The value for the constant k was calculated with the script setk.run (k = 3.046875). In TNT the "New Technology Search" option was used following Hermes et al. (2014)Hermes, M.G., Melo, G.A.R., Carpenter, J.M., 2014. The higher level phylogenetic relationships of the Eumeninae (Insecta, Hymenoptera Vespidae), with emphasis on Eumenes sensu lato. Cladistics 30, 453-484. and Hermes and Oliveira (2016)Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200..
The visualization and editing of the cladograms, as well as optimization of the characters, were performed in Winclada, with only unambiguous changes shown. Support for the branches was investigated through Symmetric Re-sampling (Goloboff et al., 2003Goloboff, P.A., Farris, J.S., Källersjö, M., Oxelmann, B., Ramírez, M., Szumik, C., 2003. Improvements to resampling measures of group support. Cladistics 19, 324-332.) using TNT 1.5, with 1000 replications for traditional (TBR) tree search and 10,000 re-sampling replications.
Results and discussion
The morphological character list is shown in Table 1. The phylogenetic analysis with implied character weighing of the matrix (Table 2) containing 42 characters returned one most-parsimonious tree (Fig. 1), with length (L) = 100, consistency index (CI) = 49, and retention index (RI) = 72. However, the resampling analysis presented small changes when compared to the implied weighting tree. These changes are related to the placement of M. tricoloratum (Zavattari, 1911), M. mexicanum (de Saussure, 1857) and M. brethesi (Bertoni, 1927) (Fig. 1).
Morphological character list for the cladistic analyses of Alphamenes. Figures that do not appear in the text are available in Supplementary Material.
Character matrix used for phylogenetic analyzes. The symbols ‘–’ and ‘?’ correspond to inapplicable and unobserved data, respectively.
Most parsimonious tree obtained with implied weighing of the characters (k = 3.046875), with length (L) = 100; consistency index (CI) = 49; retention index (RI) = 72. Black rectangles correspond to unique transformations (synapomorphies) and white rectangles correspond to homoplastic transformations. Support values (symmetric resampling) are given below branches (GC values) (where values are missing, branches collapsed during the resampling analysis). Only unambiguous changes are shown.
The generic relationships recovered herein are identical to the hypothesis presented by Hermes et al. (2014)Hermes, M.G., Melo, G.A.R., Carpenter, J.M., 2014. The higher level phylogenetic relationships of the Eumeninae (Insecta, Hymenoptera Vespidae), with emphasis on Eumenes sensu lato. Cladistics 30, 453-484.. However, only one synapomorphy (vs. five synapomorphies recovered by Hermes and Oliveira (2016)Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.) and two homoplasies (vs. one recovered by Hermes and Oliveira (2016)Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.) support Minixi as a natural group. Of these, the interantennal region raised and rounded (char 5[0]) and the lateral surface of pronotum depressed (char 11[0]) were recovered in both studies. It is worthy of mentioning the fact that the relationships among species included in Minixi changed from our previous results (see Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.). The hypothesis recovered herein allows the recognition of two major clades, which would also allow their treatment as separate genera (for example, returning to the concept of Minixi and Pachyminixi of Giordani Soika (1978)Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420., with the inclusion of M. mariachii and M. atrum in the latter). Nevertheless, we advocate the concept of Hermes and Oliveira (2016)Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200. and recognize only one genus, based on the low resampling support for the M. atrum–M. bifasciatum (von Shculthess, 1904) clade (Fig. 1) and the transitional morphological condition of M. mariachiiHermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200. (see Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200. for details).
Alphamenes was recovered as monophyletic with a somewhat high support value. Also, this is the first study to include all recognized species within the taxon in a phylogenetic investigation. Two synapomorphies related to male genitalia support the clade: basal plate of aedeagus short (char 40 [0]) and digitus with median projection adjacent to lamella (char 41 [1]) (Figs. 8–15). The male genitalia has already been pointed as providing strong evidence for the monophyly of Alphamenes (Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.; Oliveira et al., 2017Oliveira, L.A., Ferreira, W.D., Inez, T.G., Hermes, M.G., 2017. Contributions to the taxonomy of Minixi Giordani Soika and Alphamenes van der Vecht (Hymenoptera, Vespidae Eumeninae), with an identification key to the species of the latter. Zootaxa 4317, 530-540.). Furthermore, features associated with the male last visible sternum (Figs. 2–7) are taxonomically important and reliable (Giordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420.). On the other hand, identification to species level of females of Alphamenes is quite difficult, and only recently Oliveira et al. (2017)Oliveira, L.A., Ferreira, W.D., Inez, T.G., Hermes, M.G., 2017. Contributions to the taxonomy of Minixi Giordani Soika and Alphamenes van der Vecht (Hymenoptera, Vespidae Eumeninae), with an identification key to the species of the latter. Zootaxa 4317, 530-540. provided further morphological evidence to distinguish some species within the genus.
Male sternum 7 (arrows): 2. Alphamenes convexus (Fox, 1899); 3. A. semiplanusGiordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420.; 4. A. insignis (Fox, 1899); 5. A. campanulatus (Fabricius, 1804); 6. A. incertus (de Saussure, 1875); 7. A. usitatus (Fox, 1899). Figs. 2, 3, 4, 5 and 7 modified from Oliveira et al., 2017Oliveira, L.A., Ferreira, W.D., Inez, T.G., Hermes, M.G., 2017. Contributions to the taxonomy of Minixi Giordani Soika and Alphamenes van der Vecht (Hymenoptera, Vespidae Eumeninae), with an identification key to the species of the latter. Zootaxa 4317, 530-540.. Scale bars for Figs. 2, 3, 4, 5 = 1 mm; Figs. 6, 7 = 0.5 mm.
Male genitalia. Figs. 8 and 9. A. campanulatus (Fabricius, 1804): 8. Gonocoxite and volsela in lateral view; 9. Aedeagus in lateral view (a. digitus; b. cuspis; c. ventral lobi of the penis valves; d. ventral marg in of aedeagus; e. acessory lobe; f. basal apodeme of aedeagus;) (modified from Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.). Figs. 10–15. Ventral lobi of the penis valves (aedeagus): 10. A. convexus (Fox, 1899); 11. A. semiplanusGiordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420.; 12. A. incertus (de Saussure, 1875); 13. A. usitatus (Fox, 1899); 14. A. richardsiGiordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420.; 15. A. insignis (Fox, 1899). Scale bars for Figs. 10–12 and 14–15 = 0.2 mm; Fig. 13 = 0.5 mm.
Within the Alphamenes, two major clades may be readily recognized. Nevertheless, these are both supported only by copulatory features, such as the shape of the last metasomal sternum's carina (Figs. 2–7) and features related to the aedeagus (Figs. 8–15). In general, few synapomorphies were recovered to support internal relationships within Alphamenes, especially because most characters were extracted from females. As mentioned previously (Giordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420.; Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.; Oliveira et al., 2017Oliveira, L.A., Ferreira, W.D., Inez, T.G., Hermes, M.G., 2017. Contributions to the taxonomy of Minixi Giordani Soika and Alphamenes van der Vecht (Hymenoptera, Vespidae Eumeninae), with an identification key to the species of the latter. Zootaxa 4317, 530-540.), female homogeneity hampers taxonomy of the genus, but is also another evidence for monophyly of the group.
Conclusion
The character matrix and the taxon sampling in the present study were increased from previous investigations, which resulted in a more robust test for the monophyly of the included taxa. This is the first contribution to include all species recognized within Alphamenes, and the monophyly of the taxon was recovered with somewhat strong support. As mentioned in previous studies, characters of male genitalia were important in recovering both the monophyly of Alphamenes as well as internal relationships among its species.
The taxonomy of Alphamenes may be regarded as satisfactorily resolved, despite some females are still difficult to differentiate (e.g. A. semiplanusGiordani Soika, 1978Giordani Soika, A., 1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararhaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Stor. Nat. Venezia 29, 1-420., A. incertus (de Saussure, 1875), A. campanulatus (Fabricius, 1804) and A. convexus (Fox, 1899)). Some efforts may allow the resolution of these matters, such as fieldwork in order to recover nests of Alphamenes for males and females may be reared (Oliveira et al., 2017Oliveira, L.A., Ferreira, W.D., Inez, T.G., Hermes, M.G., 2017. Contributions to the taxonomy of Minixi Giordani Soika and Alphamenes van der Vecht (Hymenoptera, Vespidae Eumeninae), with an identification key to the species of the latter. Zootaxa 4317, 530-540.). From there, several morphological approaches may be conducted to sexually associate individuals of a given species, such as internal anatomy of the exoskeleton, female sting, and mouthparts. Finally, an integrative approach for Alphamenes taxonomy would most certainly serve as a solid basis for robust species hypotheses within the genus.
A series of recent contributions were recently published by the authors (e.g. Ferreira et al., 2015Ferreira, W.D., Grandinete, Y.C., Lopes, R.B., Hermes, M.G., 2015. A new contribution to the knowledge of Neotropical Eumeninae (Hymenoptera Vespidae). Zootaxa 3981, 117-124., 2017Ferreira, W.D., Oliveira, L.A., Inez, T.G., Hermes, M.G., 2017. A new species of Pirhosigma Giordani Soika, 1978 (Hymenoptera, Vespidae: Eumeninae), with additional notes and a key to the species in the genus. Zootaxa 4300, 269-278., 2018Ferreira, W.D., Oliveira, L.A., Hermes, M.G., 2018. A revision of the Neotropical eumenine wasp genus Stenosigma Giordani Soika, 1978 (Hymenoptera, Vespidae). Zootaxa 4455, 499-512.; Hermes and Oliveira, 2016Hermes, M.G., Oliveira, L.A., 2016. Morphological cladistics analysis resolves the generic limits of the Neotropical potter waps genera Minixi Giordani Soika and Pachyminixi Giordani Soika (Hymenoptera: Vespidae: Eumeninae). Invertebr. Syst. 30, 187-200.; Oliveira et al., 2017Oliveira, L.A., Ferreira, W.D., Inez, T.G., Hermes, M.G., 2017. Contributions to the taxonomy of Minixi Giordani Soika and Alphamenes van der Vecht (Hymenoptera, Vespidae Eumeninae), with an identification key to the species of the latter. Zootaxa 4317, 530-540.) regarding Neotropical eumenine taxonomy and phylogeny. The continuing practice of investigating the generic limits of Neotropical taxa using phylogenetic tools is a desirable one, especially after the expansion of generic names conducted by Giordani Soika between the 1970s and 1990s. Examples of obscure and/or dubious recognition of genera are Ancistroceroides de Saussure, 1855, Hypancistrocerus de Saussure, 1855, Hypodynerus de Saussure, 1855 (especially the eastern Neotropical fauna), Parancistrocerus Bequaert, 1925, and Stenodynerus de Saussure, 1863. Phylogenetic investigations followed by modern taxonomic revisions are desired for these taxa, for the somewhat simple task of describing new species be conducted with clear concepts about generic limits within the Neotropical eumenines.
Acknowledgements
We thank the curators cited in the Material and Methods section for the loaning of the specimens. Thanks to Dr. Salvador Arias for providing the setk.run script and the Willi Hennig Society for allowing the use of the software TNT. LAO and WDF are supported by FAPEMIG and CNPq scholarships, respectively. Financial support was provided by FAPEMIG Process CRA-APQ-02784-17.
Appendix A Supplementary data
Supplementary data associated with this article can be found, in the online version, at doi:10.1016/j.rbe.2018.11.006.
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Publication Dates
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Publication in this collection
Jan-Mar 2019
History
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Received
24 Oct 2018 -
Accepted
27 Nov 2018