Research Article |
Corresponding author: Dimitri Forero ( idf2@cornell.edu ) Academic editor: Alfred Wheeler
© 2018 Dimitri Forero, Juanita Rodríguez, Valentina Ocampo.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Forero D, Rodríguez J, Ocampo V (2018) A new species of Carvalhomiris from Colombia with an assessment of its phylogenetic position (Heteroptera, Miridae, Orthotylinae). In: Wheeler Jr AG (Ed.) A Festschrift Recognizing Thomas J. Henry for a Lifetime of Contributions to Heteropteran Systematics. ZooKeys 796: 197-214. https://doi.org/10.3897/zookeys.796.22058
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Plant bugs, species of Miridae (Heteroptera), are not well known in the Neotropics, and Colombia is not an exception. Based on data from the available systematic catalog (
Andes, genitalia, Hemiptera , Neotropical region, Zanchius group
Miridae are the most diversified family of Heteroptera with more than 11,000 described species (
Among mirid subfamilies, the Orthotylinae comprise six tribes (
Carvalhomiris is characterized by an overall pale yellow coloration with greenish areas, usually strong brachypterism in both sexes, and eyes removed from the anterior margin of the pronotum (
Recent fieldwork in Jardín, Antioquia, Colombia, led to the discovery of an additional undescribed species of Carvalhomiris. This new species represents the northernmost distribution of the genus and highlights the wide distributional gap among the known species. Additionally, the discovery of this new species allows the putative synapomorphies for Carvalhomiris proposed by
Specimens were collected near Jardín, Antioquia, Colombia, by beating vegetation along the road. Specimens were mounted, labelled, and deposited in the Entomological collection of the Museo Javeriano de Historia Natural Lorenzo Uribe S.J., of the Pontificia Universidad Javeriana, Bogotá, Colombia (
Genitalia were dissected and examined using a Zeiss Discovery V20 stereoscope or a Nikon SMZ1270 stereoscope following
Terminology for male genitalia follows
In proposing a rationale and terminology for the endosomal sclerotizations of male Austromirini,
Based on the taxonomic revision of Carvalhomiris (
For the phylogenetic analyses, we used parsimony as the optimality criterion. With the number of terminals very small, an exact solution for the matrix can be provided using a branch-and-bound algorithm (
COLOMBIA, Antioquia, Jardín, Alto de Ventanas, carretera a Riosucio, 05.5400833°N 75.8035167°W, 2913m, 2 January 2014, D. Forero.
Holotype male, pinned dry brachypterous specimen (figure 1). Original printed labels: “COLOMBIA, Antioquia, Jardín, Alto de Ventanas, carretera a Riosucio, 05.5400833°N 75.8035167°W, 2913m, 2 Ene 2014, D.Forero” / “ex. Acmella sp. (Compositae)” / “
2♂ macropters, same data as holotype /
1 nymph, same data as holotype /
Recognized by the large, basally constricted process on posterior margin of right paramere in males.
Brachypterous male. Coloration (Figure
Carvalhomiris henryi sp. n. Male genitalia. A Right paramere, lateral right view B–D left paramere B caudal view C dorsal view D medial view. Arrows on C and D show the dorsal process E, F genital capsule E caudal view F lateral left view G, H aedeagus, not everted G lateral right view H dorsal view.
Macropterous male (Figure
Brachypterous female (Figure
Macropterous female (Figure
Carvalhomiris henryi sp. n. Female genitalia. A Dorsal labiate plate and sclerotized rings, dorsal view B anterior wall, anterior view, showing the vestibulum, indicating the asymmetrical medial margin of gonapophysis 8 C posterior wall, dorsal view, showing the interramal dorsal processes.
Measurements of C. henryi sp. n. Abbreviations are as follows. Ant1: antennal segment 1. Brach: brachypter. Cun: cuneus. Cun-clyp: cuneus-clypeus. IntOcDi: interocular distance. Macr: macropter. Mesoscut: mesoscutellum. Pron: pronotum. Scut: scutellum. NA: not available.
Measurements | Length | Width | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
Total | Cun-Clyp | Head | Pron | Mesoscut | Scut | Cun | Head | Pron | IntOcDi | Ant1 | ||
Male brach | Mean | 2.96 | 2.50 | 0.44 | 0.55 | NA | 0.38 | NA | 0.66 | 0.72 | 0.38 | 0.11 |
SD | 0.07 | 0.07 | 0.05 | 0.02 | NA | 0.03 | NA | 0.03 | 0.04 | 0.02 | 0.02 | |
Range | 0.18 | 0.18 | 0.15 | 0.06 | NA | 0.06 | NA | 0.07 | 0.11 | 0.04 | 0.04 | |
Min. | 2.85 | 2.39 | 0.36 | 0.53 | NA | 0.34 | NA | 0.61 | 0.65 | 0.35 | 0.09 | |
Max | 3.03 | 2.57 | 0.51 | 0.59 | NA | 0.40 | NA | 0.68 | 0.76 | 0.39 | 0.13 | |
Female brach | Mean | 2.47 | 1.85 | 0.35 | 0.38 | NA | 0.35 | NA | 0.54 | 0.62 | 0.30 | 0.07 |
SD | 0.04 | 0.10 | 0.03 | 0.01 | NA | 0.16 | NA | 0.01 | 0.06 | 0.00 | 0.01 | |
Range | 0.05 | 0.14 | 0.04 | 0.02 | NA | 0.23 | NA | 0.01 | 0.08 | 0.00 | 0.01 | |
Min. | 2.44 | 1.78 | 0.33 | 0.37 | NA | 0.23 | NA | 0.53 | 0.58 | 0.30 | 0.06 | |
Max | 2.49 | 1.92 | 0.37 | 0.39 | NA | 0.46 | NA | 0.54 | 0.66 | 0.30 | 0.07 | |
Male macr | Mean | 4.51 | 3.73 | 0.30 | 0.48 | 0.30 | 0.37 | 0.78 | 0.65 | 1.06 | 0.37 | 0.09 |
SD | 0.01 | 0.05 | 0.08 | 0.10 | 0.09 | 0.02 | 0.02 | 0.01 | 0.01 | 0.04 | 0.01 | |
Range | 0.02 | 0.07 | 0.12 | 0.14 | 0.13 | 0.03 | 0.03 | 0.01 | 0.02 | 0.05 | 0.01 | |
Min. | 4.50 | 3.69 | 0.24 | 0.41 | 0.23 | 0.35 | 0.76 | 0.64 | 1.05 | 0.34 | 0.08 | |
Max | 4.52 | 3.76 | 0.36 | 0.55 | 0.36 | 0.38 | 0.79 | 0.65 | 1.07 | 0.39 | 0.09 | |
Female macr | Mean | 4.50 | 3.73 | 0.41 | 0.53 | 0.26 | 0.36 | 0.74 | 0.63 | 1.06 | 0.38 | 0.09 |
SD | 0.26 | 0.29 | 0.07 | 0.01 | 0.00 | 0.03 | 0.06 | 0.00 | 0.07 | 0.02 | 0.04 | |
Range | 0.37 | 0.41 | 0.10 | 0.02 | 0.00 | 0.04 | 0.08 | 0.00 | 1.04 | 0.37 | 0.07 | |
Min. | 4.31 | 3.52 | 0.36 | 0.52 | 0.26 | 0.34 | 0.70 | 0.63 | 0.07 | 0.02 | 0.04 | |
Max | 4.68 | 3.93 | 0.46 | 0.54 | 0.26 | 0.38 | 0.78 | 0.63 | 1.11 | 0.39 | 0.11 |
We are pleased to dedicate this new species to our friend and colleague Thomas J. Henry. The new name is treated as a Latin noun in the genitive case. Tom is an indefatigable entomologist, always willing to share his knowledge and help others. Tom’s enthusiasm for Heteroptera is contagious, not only in the lab but also in the field. Tom collected a long series of specimens of Carvalhomiris in Ecuador in 1996 that was described as C. bifurcatus (
Known from the type locality on a road near Jardín, Antioquia, in the western Cordillera in Colombia. It was found from 2640 to 2913 meters in elevation, in a high Andean cloud forest (Figure
Specimens of the new species were mostly associated with small roadside herbs of Acmella sp. (Asteraceae) (Figure
Species of Carvalhomiris are rather homogeneous in external morphology, but their male genitalia are distinct (
Heteropteran distributions have been little documented in Colombia, especially for geographically restricted taxa such as species of Carvalhomiris in the high Andes. An increased knowledge of biogeographic patterns might reveal diversification processes that produced these distributional ranges. Unknown is what is driving speciation processes in the Andes, a question that relates to the natural history of Carvalhomiris. It is not certain that species of this genus are strictly phytophagous. They might be omnivores, as is the case in certain orthotylines (
The collection of C. henryi sp. n. from a composite, Acmella sp., represents the second plant association for a species of Carvalhomiris. Feeding habits of the new species are unknown, but it might not be strictly phytophagous. The family Tropaeolaceae, on which C. truncatus has been found (
1 | Right paramere with ventral portion greatly elongated and apically bifurcated; brachypterous male and female with apex of hemelytron acute; medial margin of first gonapophysis nearly symmetrical | bifurcatus Forero, 2009 |
– R | ght paramere neither greatly elongated nor strongly bifurcated on ventral portion; brachypterous male and female with apex of hemelytron rounded; anterior medial margin of first gonapophysis strongly asymmetrical | 2 |
2 | Posterior serrated margin of right paramere nearly straight | brachypterus Maldonado & Ferreira, 1972 |
– P | sterior serrated margin of right paramere with a median process | 3 |
3 | Median process of posterior margin of right paramere large, nearly truncate, with wide base; dorsal portion of endosomal spicule with apex strongly left-curved; dorsal process of left paramere short, base wide | truncatus Forero, 2009 |
– M | dian process of posterior margin of right paramere small, constricted basally (Fig. |
henryi sp. n. |
The implicit enumeration search strategy for the phylogenetic analysis resulted in two equally parsimonious trees (Figure
The monophyly of Carvalhomiris was suggested without the testing of a cladistic hypothesis, but a few putative synapomorphies were noted (
Further collecting in the wide geographic gap between the known localities of Carvalhomiris species might reveal additional species and allow further testing of phylogenetic relationships in the genus.
We thank Douglas Knapp (Reserva la Esperanza, Jardín), who provided logistical support in the field. Mateo Hernández (Sopó, Cundinamarca) kindly helped us identify the plants. Financial support was provided to JRS and VO from the Centro de Fomento de la Identidad y Construcción de la Comunidad, Vicerrectoria del Medio, and from Decanatura de la Facultad de Ciencias de la Pontificia Universidad Javeriana. Al Wheeler kindly provided criticism and suggestions in an earlier version of the manuscript. This paper is a contribution of the project “Actividades docentes y de investigación como apoyo al conocimiento de la biodiversidad colombiana” ID PPTA 00006416 of the Pontificia Universidad Javeriana.
Morphological character list and their character states used in the phylogenetic analysis of Carvalhomiris species.
0. Relative body size: small (0); medium (1).
1. Antenna, type of setae on segment II medially on basal third: suberect (0); decumbent (1).
2. Antenna, length of setae on segment II medially on basal third: short (0); very short (1).
3. Antennal segment II, coloration: dark brown, pale median band (0); apical half brown, basal half pale (1); dark brown (2).
4. Antennal segment III, coloration: brown, base pale (0); brown, basal half pale yellow (1); brown, basal third pale (2); dark brown (3).
5. Pronotum, anterior lobe, structure of surface: slightly convex (0); flat (1).
6. Hemelytron development: strong brachypterism in both sexes (0); macropterous in both sexes (1).
7. Hemelytron in brachypterous specimen, relative length with respect to abdomen: short, not reaching genital capsule (0); long, reaching genital capsule (1).
8. Hemelytron in brachypterous specimen, shape of apex: acute (0); rounded (1).
9. Hemelytron, density of setae on apex (or cuneus): sparse (0); abundant (1).
10. Genital capsule, relative size to abdomen length: about half abdomen (0); about 2/3 abdomen (1); about 1/3 abdomen (2).
11. Genital capsule, structure of cephalic lateral margin in lateral view: strongly curved (0); straight (1); slightly curved (2).
12. Genital capsule, extent of ventral posterior margin relative to insertion of parameres: not projected (0); greatly projected (1).
13. Right paramere, structure of distal half: clubbed (0); nearly cylindrical (1).
14. Right paramere, structure apex: without a differentiated ventral portion (0); with a differentiated ventral portion (1).
15. Right paramere, structure of ventral portion: strongly curved dorsally (1); straight, directed caudad (1); straight, directed ventrad (2).
16. Right paramere, relative length of ventral portion: as long as apical margin (0); about half as long as apical margin (1).
17. Right paramere, structure of apical ventral portion: with a preapical spine (0); strongly bifurcated (1); as an acute process (2).
18. Right paramere, ornamentation of apical margin: gently serrate (0); strongly serrate (1); smooth, not serrate (2).
19. Right paramere, orientation of apical margin: vertical (0); reclined (1).
20. Right paramere, structure of apical margin: entire, uninterrupted (0); interrupted (1).
21. Right paramere, structure of process on apical margin interrupting margin: medium-sized spine near dorsal portion (0); large, median truncate process (1).
22. Right paramere, base of process on apical margin: constricted (0); not constricted (1).
23. Left paramere, structure of area caudad to dorsal sensory area: flat (0); with a dorsal process (1).
24. Left paramere, relative size of base of dorsal process caudad to dorsal sensory area: narrow (0); broad (1).
25. Left paramere, structure of apex: deeply cleft (0); not cleft (1).
26. Left paramere, structure on ventral surface: flat, not produced (0); with a ventral process (1).
27. Left paramere, size of apical process on ventral surface: large (0); small (1).
28. Left paramere, structure of apical process on ventral surface: acute (0); rounded (1).
29. Left paramere, structure of preapical ventral area: protruding (0); nearly flat (1).
30. Left paramere, cuticle on preapical ventral area: with medium-sized trichia (0); with very small trichia (1); flat, without trichia (2).
31. Phallotheca, structure of basal lateral area with flaplike protuberances: flat (0); with paired protuberances (1).
32. Phallotheca, structure of preapical margin: acute (0); broadly rounded (1).
33. Endosoma, number of spicules: one (0); two (1); three (2).
34. Endosomal spicule DES2, shape of spicule in lateral view: C-shaped (0); straight (1).
35. Endosomal spicule DES2, structure of the base: wide (0); narrow (1).
36. Endosomal spicule DES2, relative position in repose with respect to cephalad portion of aedeagus: close to cephalad portion (0); barely reaching caudal portion of articulatory apparatus (1).
37. Endosomal spicule DES2, C-shaped, dorsal portion, length with respect to ventral portion: three fourths of the length (0); half the length (1).
38. Endosomal spicule DES2, C-shaped, ventral portion, structure of apex: acute (0); expanded (1).
39. Endosomal spicule DES2, C-shaped, ventral portion, structure of apex: straight (0); curved dorsally (1); curved cephalad (2).
40. Endosomal spicule DES2, relative length with respect to apex of phallotheca: not reaching apex (0); reaching or surpassing apex (1).
41. First gonapophysis, symmetry of anterior medial margin: nearly symmetrical (0); strongly asymmetrical (1).
42. Dorsal labiate plate, structure of area of insertion of accessory gland: produced ventrally as a conical depression (0); flat (1).
43. Sclerotized rings of dorsal labiate plate, lateral structure: with lateral infoldings (0); without lateral infoldings (1).
44. Sclerotized rings of dorsal labiate plate, lateral infoldings structure of caudal margin: with acute process (0); rounded (1).
Matrix of morphological characters and their states used in the phylogenetic analysis of Carvalhomiris species ("–": inapplicable; "?": unknown).
Taxon | Characters | ||||||||||||||||||||||
0 | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | |
Orthotylus sp. | 1 | 1 | 0 | 2 | 3 | – | 1 | – | – | 0 | 1 | 2 | 1 | 1 | 0 | – | – | – | – | – | – | – | – |
Itacoris sp. | 0 | 0 | 0 | 0 | 2 | 1 | 1 | – | – | 0 | 2 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 2 | 1 | 0 | – | – |
Parachius sp. | 0 | 1 | 0 | 2 | 3 | 1 | 1 | – | – | 1 | 2 | 1 | 0 | 0 | 1 | 2 | 1 | 2 | 2 | 0 | 0 | – | – |
C. bifurcatus | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | – | – |
C. brachypterus | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | – |
C. truncatus | 0 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 |
C. henryi sp. n. | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 2 | 0 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 |
23 | 24 | 25 | 26 | 27 | 28 | 29 | 30 | 31 | 32 | 33 | 34 | 35 | 36 | 37 | 38 | 39 | 40 | 41 | 42 | 43 | 44 | ||
Orthotylus sp. | 0 | – | 0 | 0 | – | – | 1 | 2 | 2 | 1 | 2 | 1 | 1 | 1 | – | – | – | 1 | 0 | 1 | 0 | 1 | |
Itacoris sp. | 0 | – | 1 | 0 | – | – | 1 | 2 | 0 | 1 | 1 | 1 | 1 | 1 | – | – | – | 1 | ? | ? | ? | ? | |
Parachius sp. | 0 | – | 1 | 1 | 1 | 1 | 1 | 2 | 0 | 1 | 0 | 1 | 1 | 1 | – | – | – | 1 | 0 | 1 | 0 | 1 | |
C. bifurcatus | 0 | – | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | |
C. brachypterus | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 0 | – | |
C. truncatus | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 0 | – | |
C. henryi sp. n. | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 1 | 0 | 0 | – |
Carvalhomiris matrix (phylogenetic)