Minagrion caldense, Santos (1965b) Minagrion franciscoi Minagrion franciscoi Machado & Bedê (2015) : 288 Revision of the genus Minagrion Santos, 1965 (Odonata: Coenagrionidae) Vilela, Diogo Silva Anjos-Santos, Danielle Koroiva, Ricardo Cordero-Rivera, Adolfo Guillermo-Ferreira, Rhainer Zootaxa 2020 2020-06-02 4786 2 176 198 43J5J Santos, 1965 , Santos 1965 [151,557,825,852] Insecta Coenagrionidae Minagrion Animalia Odonata 4 180 Arthropoda species caldense   Figs. 1a–b ( ♂hab), 2a–d ( ♂and ♀tub), 3a–h ( ♂app), 4a−b ( ♂lig), 5a–g ( ♀hab), 6a–b ( ♀lob), 7a (ptx), 8a–b ( ♂app, lost holotype), 9a–c ( ♀lost allotype), 10a–c (alive ♂and ♀♀, respectively)    Minagrion caldense, Santos (1965b): 8−12 (original description ♂and ♀);— Lencioni (2006): 169 (illustrated guide; distribu- tion);—Garrison e t al.(2010): 287 (synonymic list);— Machado & Bedê (2015): 290−291, 294 (distr., illustr., comparison with  Minagrion franciscoi).     Minagrion franciscoi Machado & Bedê (2015): 288−291 (original description ♂). New synonymy.    Holotype ♂and allotype ♀: BRAZIL, Minas Gerais State, Poços de Caldas, Campo do Aterrado,  05.xii.1964, N.D. Santos& J. Machadoleg., in MNRJ[examined]  Specimens examined.  Total: 30♂♂, 20♀♀. 6♂♂, 4♀♀ BRAZIL, Minas Gerais State, Poços de Caldas, Campo do Aterrado,  23.x.1964, O. Roppa& O. Leoncinileg.,  1♂, 2♀♀in ABMM, 1♂in FAALand 4♂♂, 2♀♀in MNRJ; Minas Gerais State, São Roque de Minas, Serra da Canastra National Park( 20°14’37” S, 46°26’47” W) in different dates and collections as follows: 3♂♂,  09.x.2014 (including the holotypeand paratypesof  M. franciscoi), L.C. Bedêleg., in ABMM; 1♂,  25.ii.2015, L.C. Bedêleg., in ABMM; 4♂♂, 9♀♀,  08.iii.2018, R. Guillermo−Ferreiraleg. (of those, 2♂♂, 2♀♀in LESTES, 2♂♂, 7♀♀in ECOEVO);  3♂♂, 1♀,  19.x.2017, R. Guillermo−Ferreiraleg., in LESTES; 3♂♂,  20.x.2017, R. Guillermo−Ferreiraleg., in LESTES; 2♂♂,  21.x.2017, R. Guillermo−Ferreiraleg., in LESTES; 2♂♂, 2♀♀, Minas Gerais State, São Roque de Minas, Serra da Canastra National Park, Ponte1 (20°14’25”, 46°35’15”),  21.x.2017, D.S. Vilelaleg.   Additions to the original female description. Head( Fig. 5a–b, d–e). Dorsally black except for the occipital bar: yellow in juvenile females, pale/white in mature females; from labrum to postfrons: yellow in juvenile females, pale/blue in mature females.  Thorax( Figs. 5a–g, 6a–b, 7a, 9a). With a thick black humeral stripe extending from prothorax to posterior margin of middorsal carina; some females with lateral black stripe on the upper half of mesepimeron; remainder areas yellow in juvenile females and blue/yellow in mature females, remainder of mesepisternum, pale/white; posterior lobe of prothorax slightly convex, directed posteriorly; posterior margin of prothorax slightly widens towards the base in lateral view.  Legs( Fig. 5b, e). Pale yellow with darker areas on femoral armature in juvenile females; legs white with a black line on femoral armature in mature females.  Wings( Fig. 5b, e, g). Hyaline with a light brown pterostigma, occupying one cell; 8−9 Px in HW, 9 Px in FW; CuA extending for 8 cells distal to vein descending from subnodus in FW and 7 inHW; MP not reaching wing posterior margin and merged to MP&AA in FW and HW.  Abdomen( Figs. 5b–c, e–f, g, 9b–c). Juvenile females: S1−5 yellow with slightly darker areas laterally; S6−8 al- most orange with darker areas laterally; S9−10 yellow/orange, cercus yellow. Mature females: S1−3 white laterally, black dorsally; S4 light blue laterally, black dorsally; S5−7 blue, interrupted by thin black dorsal stripes and black basal rings; S8−10 black, with white pruinosity on older females; cercus white. On all females, notopleural suture a smoothly curved line; anterior portion of tubercle ventrally pronounced, apex slightly excavated; genital valves surpassing S10 margin for a distance as long as the segment. Measurements. Total 28−32; abdomen 22−26; FW 18−20; HW 14−17.   Diagnosis.Males of  M. caldensecan easily be separated from congeners by the almost unique excavated morphology of tubercle ( Fig. 2a–b). Females can be separated from females of other species by the following character combination: posterior lobe of prothorax with no separated lobes, middle lobe convex (concave in  M. canaanense) ( Fig. 6a–b); tubercle with a minor excavation, anterior portion ventrally pronounced ( Fig. 2c–d).   FIGURE 1.Male habitus: (a)  caldense, São Roque de Minas, at ECOEVO; (b)  caldense, Poços de Caldas, at FAAL; (c)  canaanense, Santa Tereza, at FAAL; (d)  mecistogastrum, Salesópolis, at FAAL; (e)  ribeiroi, Recreio dos BandeirantesPARATYPE, at FAAL; (f)  waltheri, São Gonçalo do Rio Preto, at ABMM.  Molecular analysis.The target COI sequences of all samples were amplified and sequenced successfully. Sequences of old specimens of  M. caldensewere also analyzed, however, they were highly fragmented and therefore could not be used in this study. The analyzed males and females of  M. caldenseformed a monophyletic group with strong support with the six females analyzed (Bayesian posterior probability, BPP = 1.00) and formed a unique clade with no further divided branches (full tree available in Appendix A). The K2P distance of COI sequences among the  M. caldensefemale typeswas 0.00% − 0.20%, while for the other species this value ranged between 13.20% and 20.10% (see Appendix B). These evidences implied that the males and females analyzed here belong to the same species.  Ontogenetic color change.The dissections made in both female stages of  M. franciscoilead us to the conclusion that females of this species undergo a complete ontogenetic coloration change, therefore representing juvenile (yellow females) and mature (blue females) instead of two different morphs. Juvenile females only presented undeveloped egg masses and all examined mature females presented fully developed eggs with an oval shape and resilient chorion ( Waage 1980, Higashi & Watanabe 1993).   Remarks.Upon the examination of holotypesand paratypesof  M. caldenseof  M. franciscoi, we found no substantial characters in morphology to separate them, although some traits such as cercus ( Figs. 3a–h, 8a–b) and genital ligula ( Fig. 4a–b) present subtle differences, which can be interpreted as intraspecific variation. Machado & Bedê (2015: 291)state that the sclerotized lateral process in genital ligula is separated, forming two structures on each side in  M. franciscoi. We noticed a subtle variation (size, shape) of this trait on the examined males, but we saw no such separation among material (including paratypes) identified as this species. The sclerotized lateral processes in all were entire (as in Fig. 4a–b). Excluding this character, coloration was the diagnostic means with which to separate  M. caldensefrom  M. franciscoiby Machado & Bedê (2015). However, this can be problematic since coloration has proven not always to be a reliable source of taxonomic diagnosis in Coenagrionids (Vilela et al.2019). In both ‘species’, the posterior lobe of female prothorax is slightly convex, posteriorly oriented, and the lateral margins of prothorax slightly widening towards the base, in lateral view. Females of  M. caldenseare also morphologically identical to females that we associated with males of  M. franciscoi. As in males, coloration accounted for the main differences between these females which cannot be used as a specific differential since females undergo ontogenetic changes in coloration. Therefore we suggest that  M. franciscoibe treated as a junior synonym of  M. caldense.   Distribution.This species remains one of the rarest in the genus. The only two known localities of occurrence are in Poços de Caldas ( typelocality), and the National Park of Serra da Canastra both in Minas Gerais State. 1964-12-05 MNRJ N. D. Santos & J. Machado Brazil Campo do Aterrado Pocos de Caldas 4 180 2 1 1 Minas Gerais holotype 1964-10-23 O. Roppa & O. Leoncini Brazil Total Campo do Aterrado Pocos de Caldas 4 180 60 24 36 Minas Gerais 2014-10-09 ABMM, FAAL, MNRJ L. C. Bede Brazil -20.243612 Serra da Canastra National Park 21 -46.44639 Sao Roque de Minas 4 180 13 4 9 Minas Gerais holotype [274,755,1343,1370] 2015-02-25 ABMM L. C. Bede Brazil 4 180 1 1 Minas Gerais holotype 2018-03-08 ABMM, R, ECOEVO Guillermo-Ferreira Brazil 4 180 26 18 8 Minas Gerais holotype [885,1432,1380,1407] 2017-10-19 R Guillermo-Ferreira Brazil 4 180 4 1 3 Minas Gerais holotype [151,813,1416,1443] 2017-10-20 R Guillermo-Ferreira Brazil 4 180 3 3 Minas Gerais holotype 2017-10-21 R Guillermo-Ferreira Brazil 4 180 2 2 Minas Gerais holotype 2017-10-21 D. S. Vilela Brazil Ponte Sao Roque de Minas 4 180 4 2 2 Minas Gerais holotype