A phylogeny of Carrerapyrgota Aczél (Diptera, Pyrgotidae)

Abstract

CarrerapyrgotaAczél, 1956 is a Neotropical Diptera genus within Pyrgotidae. Currently, the genus comprise four species: C. miliariaAczél, 1956 (type species), C. personata (Lutz & Lima, 1918), C. aczeliMello, Lamas & Rafael, 2010, and C. bernardiiMello, Lamas & Rafael, 2010. This study investigates the phylogenetic relationships among Carrerapyrgota species, aiming to establish their monophyly as a genus. To achieve this, we conducted phylogenetic analyses based on 22 external morphological characters of adults, emploing the principle of parsimony. Analyses using both equal and implied character weighting support the monophyly of the genus, based on the following apomorphies: absence of the postscutellum, R₂₊₃ with spurious apical vein, lower calypter with a longitudinal row of bristles, and the presence of a posterior longitudinal groove on the female fore femur. The species C. aczeli, C. bernardii, and C. miliaria form a monophyletic group, supported by the synapomorphic condition of a spot on the forehead. Within this group, C. bernardii and C. miliaria are sister species, sharing the synapomorphic condition of a less sclerotized M vein, after dm-cu compared to the anterior region, as well as the homoplastic condition of a spot on the median occipital sclerite. This phylogenetic analysis reaffirms the synonymy proposed by Bernardi (1990), which treats AnapyrgotaSteyskal, 1967, as a junior synonym of Carrerapyrgota. The evolutionary relationship among the species of the genus is represented as (C. personata (C. aczeli (C. bernardii, C. miliaria))). Furthermore, this study includes an updated distributional map for the genus, featuring the first record of Pyrgotinae in the Andean region.

Keywords:
Cladistic; Monophyly; Pyrgotinae; South America; Tephritoidea

INTRODUCTION

Pyrgotidae is a dipteran family distributed worldwide, comprising approximately 365 species across 55 genera (Korneyev, 2006). In the Neotropics, 58 species are recognize, divided into 12 genera (Steyskal, 1967; Bernardi, 1991; Mello et al., 2010; Mello & Lamas, 2014). Members of this family are nocturnal, with larvae developing as endoparasitoids of adult beetles in the Scarabaeidae family (Forbes, 1908; Davis, 1913, 1919; De Meijere, 1916; Wolcott, 1922; Aldrich, 1928; Clausen et al., 1933; Moutia, 1940; Ritcher, 1940; Gardner & Parker, 1940; Paramonov, 1958).

Carrerapyrgota was proposed by Aczél (1956a) to host the new species Carrerapyrgota miliaria, described based on male and female specimens from Brazil and Argentina. Lutz & Lima (1918) described the species Apyrgota personata based on a single specimen (of unidentified sex, due to the lack of the abdomen) from Pernambuco, Brazil. Hennig (1936) and Aczél (1956b) noted the absence of the genus Apyrgota in the neotropics, and suggested that the species described by Lutz & Lima should be transferred to the genus Pyrgota (Hennig, 1936) or included in a new genus (Aczél, 1956b). Steyskal (1967) presented a taxonomic catalog of Neotropical Pyrgotidae, proposing the genus Anapyrgota to host the species A. personata.Bernardi (1990) considered Anapyrgota synonymous with Carrerapyrgota, proposing C. personata as a new combination with Carrerapyrgota.

Mello et al. (2010) provided a revision of Carrerapyrgota, which included the description of two new Brazilian species, C. aczeliMello, Lamas & Rafael, 2010 and C. bernardiiMello, Lamas & Rafael, 2010. They retained the synonymy and new combination proposed by Bernardi (1990), reported the first South American pyrgotid species of pyrgotids associated with a host (one beetle species), and presented an identification key and a distributional map of the species.

Carrerapyrgota is defined by the following combination of characters: medial vertical seta intersecting with postocellar seta; mesofacial plate without carina; one notopleural seta; postscutellum absent; female forefemur featuring a posterior longitudinal groove; vein C with Sc break; Sc incomplete (not reaching C); R₂₊₃ with an apical spur vein; lower calypter covered by a longitudinal row of hairs; and ovipositor without apical hook.

This study aims to investigate the phylogenetic relationships among Carrerapyrgota species, including their monophyly as a genus, and to test the synonymy proposed by Bernardi (1990). To accomplish this, phylogenetic parsimony analyses were performed based on the morphological characteristics of adults. Additionally, we present the first record of a Pyrgotinae from the Andean region, for the species C. miliaria.

MATERIAL AND METHODS

Specimens used in this study were belongs from the following institutions: Coleção Entomológica “Padre Jesus Santiago Moure”, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Brazil (DZUP); Fundação e Instituto Oswaldo Cruz, Rio de Janeiro, Brazil (FIOCRUZ); Fundación e Instituto Miguel Lillo, San Miguel de Tucumán, Argentina (IMLA); Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil (INPA); Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP); e Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ). The material from MNRJ has been preserved since the 2018 fire incident (Couri, personal communication). The complete list of specimens studied here corresponds to that presented by Mello et al. (2010). Additional specimens were obtain from the collection of the Instituto Argentino de Investigaciones Zonas Áridas, Mendoza, Argentina (IADIZA).

Terminology in this study adheres to the conventions outlined by Korneyev (2006), Cumming & Wood (2009), and Mello et al. (2010). Photographs and illustrations were edited using Adobe Photoshop CC 2014, and Adobe Illustrator CS. The distribution map was crated using QGIS 3.28.2.

Outgroup taxa utilized for this analysis consisted of Ceratitis capitata (Wiedemann, 1824) (Tephritidae, Dacinae); Descoleia teretruraAczél, 1956 (Tephritoidea incertae sedis); Leptopyrgota sahlbergiana Frey, 1918 (Pyrgotidae, Pyrgotinae); Idiopyrgota setiventrisAczél, 1956 (Pyrgotidae, Pyrgotinae); and Stenopyrgota crassitibiaAczél, 1956 (Pyrgotidae, Pyrgotinae). The cladograms were rooted between C. capitata and the remaining taxa.

Phylogenetic parsimony analyses were conducted using data related to the external morphology of adult males and females. Morphological characters, whether binary or multistate, were coded and organized into a matrix using the Mesquite v3.61 software. Character codification followed the logic proposed by Sereno (2007). Multistate characters were treat as unordered (Fitch, 1971). The analysis was performed using the Tree Analysis Using New Technology v1.1 (Goloboff et al., 2008).

Analyses were carried out with equal weighting of characters using heuristic searches with 1,000 replicates, retaining up to 100 cladograms. Additionally analyses were conducted with implied weighting employing concavity K values ranging from 1 to 10. Lower K values, approaching zero, imposed stronger penalties on characters displaying homoplasy (Goloboff, 1993). This range of K values was selected to test the phylogenetic hypothesis under differential weighting. Results for each analysis were summarized through strict consensus (Sokal & Rohlf, 1981).

Bremer’s Absolute Supports were calculated for the cladograms resulting from the equal weighting analysis (Bremer, 1994), while Bremer’s Relative Support was employed for the implied weighting analysis (Goloboff & Farris, 2001). Figures of the cladograms were edited using the WINCLADA v1.00.08 software (Nixon, 2002).

RESULTS AND DISCUSSION

A total of 22 morphological characters were analyzed, including one related to the general body, nine concerning the head, 11 regarding the thorax, and one specific to the male abdomen (Table 1). The length (L), consistency index (CI), and retention index (RI) are provided in parentheses.

Table 1
Matrix of Morphological Characters used for Carrerapyrgota and outgroups.

Taxon/Character	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22
Ceratitis capitata	1	0	1	1	0	1	0	0	1	1	0	1	3	1	1	0	0	0	1	0	0	1
Descoleia teretrura	0	0	0	1	1	1	0	1	-	0	0	0	3	0	1	0	0	0	0	0	0	1
Stenopyrgota crassitiba	1	0	1	1	0	2	1	1	-	0	0	-	1	1	0	0	1	0	0	0	0	-
Idiopyrgota setiventris	0	0	0	0	-	2	1	0	2	1	0	0	2	1	0	0	1	0	0	0	0	0
Leptopyrgota sahlbergiana	0	0	1	0	1	0	1	1	0	0	0	0	0	0	1	1	1	0	0	0	0	?
C. aczeli	0	1	1	1	-	2	1	1	2	0	1	1	2	1	1	1	1	0	1	1	1	1
C. bernardii	1	1	1	0	1	2	1	1	2	1	1	1	1	1	1	1	1	1	0	1	1	1
C. miliaria	1	1	1	0	1	2	1	1	2	1	1	1	1	1	1	1	1	1	0	1	1	0
C. personata	1	0	1	0	1	2	1	1	2	0	1	1	3	1	1	1	1	0	1	1	1	-

List of Characters

1. Body setae and setulae, color (L: 3; CI: 33; RI: 33): (0) reddish yellow; (1) black.

2. Frons, spot (L: 1; CI: 100; RI: 100): (0) absent (Fig. 1); (1) present (Fig. 2).

3. Antennal groove, median carina (L: 2; CI: 50; RI: 0): (0) present; (1) absent.

4. Antennal groove, lower margin, spot (L: 2; CI: 50; RI: 66): (0) present; (1) absent.

5. Pedicel, dorsal surface, size in relation to the first flagellomere (L: 2; CI: 50; RI: 0): (0) shorter; (1) longer (Fig. 1).

6. Arista, position in the dorsal surface of the first flagellomere (L: 3; CI: 66; RI: 50): (0) apical (Fig. 1); (1) basal; (2) middle (Fig. 3).

7. Ocelli (L: 1; CI: 100; RI: 100): (0) present (Fig. 4); (1) absent (Fig. 1).

8. Ocellar seta (L: 2; CI: 50; RI: 0): (0) present; (1) absent (Fig. 1).

9. Postocellar seta, position in relation to the medial vertical seta (L: 3; CI: 66; RI: 0): (0) convergent, not crossed; (1) parallel; (2) convergent, crossed like X.

10. Median occipital sclerite, spot (L: 3; CI: 33; RI: 33): (0) absent; (1) present.

11. Postscutelum (L: 1; CI: 100; RI: 100): (0) present; (1) absent.

12. Notopleural seta (L: 2; CI: 50; RI: 50): (0) two setae; (1) one seta.

13. Wing, color pattern (L: 6; CI: 50; RI: 40): (0) hyaline with apical spot (Fig. 5); (1) reticulate (Fig. 6); (2) bicolor (Fig. 7); (3) banded (Fig. 8).

14. Costal vein (L: 2; CI: 50; RI: 0): (1) ending at vein M; (0) ending before reach vein M.

15. Costal vein (L: 2; CI: 50; RI: 0): (0) unbroken on Sc; (1) broken on Sc.

16. Vein R₂₊₃, apical spur vein (L: 1; CI: 100; RI: 100): (0) absent (Figs. 5-7); (1) present (Figs. 8-9).

17. Vein R₄₊₅, dorsal surface (L: 1; CI: 100; RI: 100): (0) setulose; (1) bare.

18. Vein M, sclerotization (L: 1; CI: 100; RI: 100): (0) uniformly esclerotized (Figs. 5-8); (1) less sclerotization in front of dm-cu (Fig. 9).

19. Vein M, in front of dm-cu, position in relation to the vein R₄₊₅ (L: 3; CI: 33; RI: 0): (0) straight (Figs. 5-7); (1) sinuose (Figs. 8-9).

20. Lower calypter, longitudinal row of hairs (L: 1; CI: 100; RI: 100): (0) absent; (1) present.

21. Female forefemur, posterior surface, longitudinal groove (L: 1; CI: 100; RI: 100): (0) absent; (1) present (Fig. 10).

22. Male sternite 5, anterior margin, shape (L: 3; CI: 66; RI: 0): (0) bilobate (Fig. 11); (1) straight (Fig. 12); (2) rounded.

Figures 1-4
Heads: dorsal view of Leptopyrgota juniae Bernardi (1); Carrerapyrgota miliaria Aczél (2); Teretrura tinctipennis Malloch (4); lateral view of Idiopyrgota setiventris Aczél (3). Scales: Fig. 1: 0.5 mm; Figs. 2-3: 1 mm; Fig. 4: 0.25 mm. Fig. 2 modified from Mello et al. (2010), Fig. 3 modified from Mello & Lamas (2008).

Figures 5-9
Wings: Leptopyrgota sahlbergiana Frey (5); Stenopyrgota crassitibia Aczél (6); Idiopyrgota setiventris Aczél (7); Carrerapyrgota personata (Lutz & Lima) (8); C. miliaria Aczél (9). Scales: Fig. 5: 2 mm; Figs. 6-9: 1 mm. Fig. 6 modified from Mello & Lamas (2014), Fig. 7 modified from Mello & Lamas (2008), Figs. 8-9 modified from Mello et al. (2010).

Figures 10-12
Female fore femur, posterior view of Carrerapyrgota aczeli Mello, Lamas & Rafael (10). Male sternite five of C. miliaria Aczél (11); C. aczeli (12). Scales: 1 mm. Figs. 10-12 modified from Mello et al. (2010).

The analysis with equal weighting yielded six most parsimonious cladograms, as indicated by the parenthetical notation in Table 2. These cladograms exhibited an L value of 43 steps, CI 62, and RI 60. In all six cladograms, the monophyly of the genus was supported by four synapomorphies: postscutellum absent (11:1), R₂₊₃ with apical spur vein (16:1. Figs. 8-9), lower calypter with a longitudinal row of hairs (20:1), and the presence of a longitudinal groove on the posterior surface of female fore femur (21:1. Fig. 10). Furthermore, all six cladograms, consistently placed C. bernardii and C. miliaria as sister groups, supported by the synapomorphic conditions of vein M being less sclerotized after dm-cu compared to its basal half (18:1. Fig. 9), as well as the homoplasic presence of a spot on the median occipital sclerite (10:1).

Table 2
Parenthetical notation of the seven cladograms most parcimonious obtained using equal and implied weighting with k values from 1-10.

Cladograms from equal weighting from implied weighting (K values)	Correspondent cladograms	Parenthetical notation
*Cladogram 1	K3, K4, K5, K6, K7, K8, and K10	((Ce. capitata (D. teretrura (S. crassitibia (I. setiventris, L. sahlbergiana)))) (Ca. personata (Ca. aczeli (Ca. bernardii, Ca. miliaria))))
Cladogram 2	((Ce. capitata (D. teretrura (S. crassitibia (I. setiventris, L. sahlbergiana)))) ((Ca. personata, Ca. aczeli), (Ca. bernardii, Ca. miliaria)))
Cladogram 3	K3, K4, K5, K6, K7, K8, and K10	((Ce. capitata (S. crassitibia (L. sahlbergiana (I. setiventris, D. teretrura)))) (Ca. personata (Ca. aczeli (Ca. bernardii, Ca. miliaria))))
Cladogram 4	K3, K4, K5, K6, K7, K8, K9, and K10	((Ce. capitata (S. crassitibia (I. setiventris (L. sahlbergiana, D. teretrura)))) (Ca. personata (Ca. aczeli (Ca. bernardii, Ca. miliaria))))
Cladogram 5	((Ce. capitata (S. crassitibia (L. sahlbergiana (I. setiventris, D. teretrura)))) ((Ca. personata, Ca. aczeli), (Ca. bernardii, Ca. miliaria)))
Cladogram 6	((Ce. capitata (S. crassitibia (I. setiventris (L. sahlbergiana, D. teretrura)))) ((Ca. personata, Ca. aczeli), (Ca. bernardii, Ca. miliaria)))
Cladogram without correspondent in equal weighting analysis K1, K2 and K3	((Ce. capitata (D. teretrura (L. sahlbergiana (I. setiventris, S. crassitibia)))) (Ca. personata (Ca. aczeli (Ca. bernardii, Ca. miliaria))))

• Number of cladogram using equal weighting: six cladograms. Number of cladogram using implied weighting with K1: 1 cladogram; K2: 1 cladogram; K3: 4 cladograms; K4-K8: three cladograms; K9: 1 cladogram; and K10: three cladograms, total of 25 cladograms. Number of cladograms using equal and implied weighting 31 cladograms, distributes in seven different hypothesis. *Cladogram selected to represents the phylogeny of Carrerapyrgota.

In three of the cladograms, the clade consisting of (C. aczeli, (C. bernardii, C. miliaria)) was supported by the presence of a spot on the frons (2:1. Fig. 2). The strict consensus of the cladograms from the equal weighting analysis, along with Bremer’s Absolute Support analysis, is presents in Fig. 13. The monophyly of the groups obtained in the equal weighting analysis, Carrerapyrgota, and the clade composed of (C. bernardii, C. miliaria), was consistently found in cladograms within two steps of the most parsimonious cladograms (Fig. 13).

Figure 13
Strict consensus cladogram and Absolute Bremer Support of Carrerapyrgota, resulted from analysis with equal weighting of characters.

Analyses with implied weighting produced a total of 25 cladograms, resulted in a single phylogenetic hypothesis for the ingroup and four distinct topologies for the outgroups (Table 2). Table 2 provides the number of cladograms obtained for each K value, along with the corresponding phylogenetic hypotheses from the equal weighting analysis.

The optimal cladogram selected to represent the monophyly of Carrerapyrgota and the phylogenetic hypothesis is decipted in Fig. 15. This cladogram corresponds to one of the most parsimonious cladograms resulting from equal weighting, and seven of the most parsimonious cladograms from analyses with implied weighting using K3-K8, and K10 (Table 2). Adittionally, this cladogram aligns with the classification of the pyrgotids proposed by McAlpine (1990), concerning the relationship of the outgroup.

Figure 14
Strict consensus cladogram and Relative Bremer Support of Carrerapyrgota, resultant from analysis with implied weighting from K3.

Figure 15
Optimal cladogram selected to represents the evolutionary relationship among the species of Carrerapyrgota. Black circles represents synapomorphies, white circles represents homoplasies.

The strict consensus, along with Bremer’s Relative Support, for the analyses with implied weighting, starting from K3, is presented in Fig. 14. The distribution of characters and their respective states pertaining to the optimal cladogram, are illustrated in Fig. 15.

New occurrences of Carrerapyrgota miliaria (Fig. 16): Argentina, Mendoza, Santa Rosa, Ñacuñán; 30/x/1996; G. Debandi col. 1 female (IADIZA), Mello & Núñez-Campero det. xi/2022. Idem, 18/vii-16/viii/1998. 1 male (IADIZA). Neuquén, Collón Curá; 21/x/1967; M. Gentile col. 1 female (IADIZA), Mello & Núñez-Campero det. xi/2022.

Figure 16
Distribution map of Carrerapyrgota species. n. oc.: new occurrences. Modified from Mello et al. (2010).

CONCLUSION

The confirmation of the monophyly of Carrerapyrgota provides further support for the synonymy proposed by Bernardi (1990), which regards Anapyrgota as a junior synonym of Carrerapyrgota. Additionally, the discovery of new occurrences of C. miliaria in Neuquén, Argentina, marks the first record of the Pyrgotinae in the Andean region. This finding expands the distributional range of the subfamily in South America.

ACKNOWLEDGMENTS:

We are very grateful to all curators of the cited collections, which made material available for our study; to the Dr. Alessandre Pereira-Colavite (UFPB), Dr. Danilo Ament (UFLA), Fernando da Silva Carvalho-Filho (MPEG), Dr. Gustavo Graciolli (UFMS), João Paulo Vinicios Rodrigues (UFPR), and to Dr. Segundo Núñez-Campero (CRILAR) for a first revision of the manuscript.

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  » https://doi.org/10.46429/jaupr.v6i1.15128

• FUNDING INFORMATION:

  Fundação de Apoio ao Desenvolvimento do Ensino, Ciência e Tecnologia do Estado de Mato Grosso do Sul (FUNDECT) (Proc. 71/700.175/2020), fellowship to the first author.

Edited by

Edited by:

Kirstern Lica Follmann Haseyama

Publication Dates

• Publication in this collection
  10 June 2024
• Date of issue
  2024

History

• Received
  03 Feb 2023
• Accepted
  01 Sept 2023
• Published
  02 Jan 2024