Research Article |
Corresponding author: Ulises Castro-Valderrama ( ucastro.11@gmail.com ) Academic editor: Yalin Zhang
© 2023 Francisco Armendáriz-Toledano, Misael Adrián López-Posadas, Youssef Utrera-Vélez, Jesús Romero Nápoles, Ulises Castro-Valderrama.
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:
Armendáriz-Toledano F, López-Posadas MA, Utrera-Vélez Y, Romero Nápoles J, Castro-Valderrama U (2023) More than 80 years without new taxa: analysis of morphological variation among members of Mexican Aeneolamia Fennah (Hemiptera, Cercopidae) support a new species in the genus. ZooKeys 1139: 71-106. https://doi.org/10.3897/zookeys.1139.85270
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The genus Aeneolamia includes eight described species and 32 subspecies widely distributed in America. In Mexico, two species (A. contigua and A. albofasciata) and one subspecies (A. contigua campecheana) are recognized. In a recent study of Cercopidae in Mexico, a new species of Aeneolamia was noted from Oaxaca, Mexico based on body color and the ornamentation patterns of tegmen, without a formal taxonomic description. To test the hypothesis of an extant new taxon within the genus a comprehensive analysis of intraspecific morphological variation from 46 morphological features was performed, four related to tegmen color patterns in both sexes, six to male genitalia, and 36 continuous characters measured in specimens of both sexes of Mexican Aeneolamia from several geographical localities using traditional univariate, multivariate morphometric, and geometric morphometric methods. This is the first time that this approach has been used in Cercopidae. Aeneolamia danpecki Castro, Armendáriz & Utrera, sp. nov. from Oaxaca showed pronounced morphological differences in tegmen coloration patterns, the shape of different elements of the male genitalia, and body measurements compared to the other Mexican members of Aeneolamia; therefore, it is described as a new species.
Aeneolamia aff. albofasciata, Cercopoidea, grasses pest, Spittlebug, sugarcane
In the Neotropical region, 60 genera of Cercopidae are integrated into the subfamily Ischnorhininae (
In the compilation of
In Mexico, Aeneolamia is represented by two species, A. albofasciata (Lallemand, 1939) and A. contigua (Walker, 1851). Both Mexican species of Aeneolamia are polyphagous on Poaceae and inhabit almost most regions from Mexican Republic (
A total of 628 Aeneolamia adults from 59 Mexican localities corresponding to 260 females and 368 males were reviewed. From the total sample, 64 specimens (43 ♀, 21 ♂) correspond to A. danpecki sp. nov., 496 to A. albofasciata (178 ♀, 318 ♂), and 68 to A. contigua (39 ♀, 29 ♂). For the third species, we included specimens collected around the respective type localities of the previously recognized subspecies A. contigua campecheana, A. contigua postica, and A. contigua sanctaerosae because the type localities were not geographically detailed in the original descriptions or the habitat of the subspecies in the locality had disappeared (Table
Species, locality, date, and sample size of Mexican Aeneolamia. The number of specimens in parentheses refer to those included in the morphometric analysis. aA. albofasciata identified by Clark in 1975 and deposited in CEAM, bA. contigua campecheana, cA. contigua postica.
Species | Locality | Date | Total | Female | Male |
---|---|---|---|---|---|
A. albofasciata | Campeche, Colegio de Postgraduados | 3/X/2016 | 199 | 79 (2) | 120 |
Campeche, Haltunchen, Km 159.5 | 2/X/2016 | 110 | 10 (1) | 100 (2) | |
Chiapas, Comunidad Providencia | 6/VI/2011 | 1 | – | 1 (1) | |
Chiapas, ECOSUR, Tapachula | 18/VI/1999 | 1 | 1 | – | |
Chiapas, Ejido Rizo de Oro, Cintalapa | 27/V/2011 | 2 | – | 2 | |
Guerrero, Acapulco | 21/VIII/1938 | 1 | 1 | – | |
Guerrero, Petaquillas, 9 km W Chilpancingo | 6/VI/1963 | 1 | – | 1 | |
Michoacán, Charapendo | 18/VIII/2015 | 1 | 1 (1) | – | |
Michoacán, Tangamandapio | 14/IX/2017 | 119 | 56 (3) | 63 (5) | |
Michoacán, Taretan | 13/IX/1963 | 2 | 2 (1) | – | |
Michoacán, Uruapán | VII/1998 | 1 | – | 1 (1) | |
Morelos, Cuatla | 14/IX/1980 | 3 | 1 (1) | 2 | |
Morelos, Cuautla, Cuautlixco | 22/V/2002 | 1 | – | 1 (1) | |
Morelos, Palo Bolero | 7/X/1995 | 1 | – | 1 (1) | |
Nuevo León, Apodaca | 4/VIII/1979 | 20 | 6 | 14 | |
Quintana Roo, Tecnológico de Chetumal | 15/X/2017 | 2 | – | 2 | |
San Luis Potosí, Sierra El Abra, Los Patos | 8/IX/2017 | 1 | 1 | – | |
Sonora, Municipio de ímuris, ímuris | 9/VIII/2013 | 1 | – | 1 | |
Sonora, Centro Invest. Pec. Est. Sonora, Carbó | X/1981 | 2 | 2 | – | |
Tabasco, Cárdenas | 28/VI/1982 | 1 | 1 (1) | – | |
Tabasco, Cárdenas | 12/I/2012 | 1 | 1 (1) | – | |
Tamaulipas, Cd. Mante | VI/1987 | 1 | 1 (1) | – | |
Tamaulipas, Cd. Mante | 7/X/1983 | 4 | 1 (1) | 3 (1) | |
a | Veracruz, Cosamaloapan | 20/VII/1962 | 1 | 1 | – |
a | Veracruz, Tecolutla | 7/IX/1973 | 1 | – | 1 |
Veracruz, Km 4.5 Carr Cardel-Salmoral | 4/IX/2003 | 14 | 12 (1) | 2 | |
Veracruz, Úrsulo Galván | 23/VI/2021 | 4 | 1 (1) | 3 (3) | |
Total | 496 | 178 (15) | 318 (15) | ||
A. contigua | Quintana Roo, 2 km S Rancho El 24 | 20/XII/1984 | 1 | – | 1 |
Chiapas, Calzada larga, Villaflores | 27/X/2012 | 1 | – | 1 | |
Chiapas, Finca Cucalhuitz, 19 Km NE Bochil | 28/XI/1961 | 1 | 1 (1) | – | |
Chiapas, Llano La Lima, Tapachula | 9/VI/2013 | 1 | 1 | – | |
Chiapas, Palenque | 31/I/1985 | 1 | 1 | ||
Chihuahua, Chihuahua | 12/VII/1938 | 1 | – | 1 | |
Guerrero, Almolonga | 30/VII/1962 | 5 | 1 (1) | 4 (3) | |
Jalisco, Ameca | 23/VII/1999 | 1 | – | 1 | |
Michoacán, Morelia | 1/VI/1963 | 1 | – | 1 (1) | |
Morelos, Cuatla | 6/II/1996 | 2 | 1 (1) | 1 (1) | |
Morelos, Reserva de la Biosfera Huahutla | 27/V/2000 | 15 | 13 (3) | 2 | |
Nayarit, Guayabitos | 2/X/1980 | 1 | 1 (1) | – | |
Oaxaca, I. Bastida | 12/IX/1981 | 1 | – | 1 (1) | |
Oaxaca, Tehuantepec | 10/VII/1966 | 1 | 1 (1) | – | |
b | Quintana Roo, Tecnológico de Chetumal | 15/X/2017 | 1 | – | 1 |
Tabasco, Cárdenas | 14/VII/1994 | 1 | – | 1 (1) | |
c | Veracruz, Cd. Mendoza | 15/IX/1994 | 2 | 1 | 1 (1) |
c | Veracruz, Colegio de Postgraduados, Campus Córdoba | 15/IX/1994 | 1 | – | 1 (1) |
Veracruz, Est. Los Tuxtlas, San Andrés | 21/IX/2007 | 1 | – | 1 (1) | |
Veracruz, Isla | 13/VII/2002 | 2 | 2 (2) | – | |
Veracruz, Km 14 Aut. Cárdenas-Minatitlán, Rancho La Majada | 4/X/2016 | 15 | 10 (2) | 5 (4) | |
c | Veracruz, La Antigua | 28/VIII/1978 | 2 | 2 (2) | – |
Veracruz, Las Vigas | 19/VI/1965 | 2 | 1 | 1 (1) | |
Veracruz, Playa Escondida, Catemaco | 13/VI/1979 | 1 | 1 (1) | – | |
Veracruz, Playa Escondida, Catemaco | 13/VI/1979 | 1 | – | 1 | |
Veracruz, Tinája | 18/IX/1994 | 1 | – | 1 | |
Yucatán, Chichen Itza | 20/VI/1985 | 2 | 1 | 1 | |
Yucatán, Ruta 295, Km 93 Rio Lagartos | 18/VI/1985 | 2 | – | 2 | |
b | Yucatán, Cuncunul | 13/IX/1994 | 1 | 1 | – |
Total | 68 | 39 (15) | 29 (15) | ||
Aeneolamia danpecki sp. nov. | Oaxaca, 5 km San Martín Lochila | 12/VII/2004 | 1 | 1 (1) | – |
Oaxaca, Sola de Vega | 28/IX/2003 | 48 | 33 (5) | 15 (9) | |
Oaxaca, La Trinidad, Zaachila | 28/VIII/2018 | 15 | 9 (9) | 6 (6) | |
Total | 64 | 43 (15) | 21 (15) | ||
TOTAL | 628 | 260 | 368 |
Taxonomic identifications of the species were based on male genitalia. In addition, we included two specimens identified as A. albofasciata (= A. albofasciata occidentalis) from
To manipulate specimens and take photographs, we used the method of Valdez-Carrasco (
Discrete morphological characters. Because of high polymorphism in color patterns of wings recorded in some species of Auchenorrhyncha families, particularly in cercopids, and due to the male genitalia traits providing robust evidence to support A. danpecki sp. nov. (
Character states to compare the variation of tegmen color patterns A, D, G, H, K, L Aeneolamia danpecki sp. nov. B, E, I, M A. albofasciata C, F, J, N A. contigua D–F dotted lines indicate the internal anterior region of tegmen. Arrows indicate the transversal lines on tegmen: G–J anterior region K–N posterior region.
Character states to compare the variation of male genitalia A, C, F, I, L, O, R Aeneolamia danpecki sp. nov. D, G, J, M, P, S A. albofasciata B, E, H, K, N, Q, T A. contigua A, B anal tube, lateral view C–E aedeagus within pygofer, lateral view F–H distal region of parameres, lateral view I–K subgenital plates, ventral view L–N subapical spines of parameres, lateral view O–Q phallobase and aedeagus, lateral view R–T phallobase and aedeagus, anterodorsal view. Abbreviations: ae aedeagus, a acuminate, as aedeagus spine, bl blunt internal distal edge, dl dorsal lobe, dp digital process of pygofer, lb lobe, pa paramere, pas primary apical spine of parameres, phb phallobase, py pygofer, rh rhomboidal apex with lateral straight edge, ss secondary subapical spine of paramere, st slight concave lateral edge, vl ventral inferior lobe, X tenth segment of anal tube, XI eleventh segment of anal tube.
Because the Aeneolamia species display apparent differences in body size, 90 adults from 31 Mexican localities were compared using measurements of the head, mouthparts, pronotum, tegmina, and legs (Fig.
Distribution and details of the continuous features used to quantify the morphological variation of Mexican Aeneolamia spp. Habitus view: A dorsal B ventral C lateral. Features Abbreviations: HWEd Head width with eyes HLd head length in dorsal view, PWd pronotum width in dorsal view PLd pronotum length in dorsal view SWd scutellum width in dorsal view SLd scutellum length in dorsal view PcLv postclypeus length in ventral view PcWv postclypeus width in ventral view Alv anteclypeus length in ventral view AWv anteclypeus width in ventral view SLv stylet length in ventral view SWv stylet width in ventral view PCWv posterior coxa width in ventral view PcLv posterior coxa length in ventral view BLWv body length without wings in ventral view PCLl postclypeus length in lateral view PCWl postclypeus width in lateral view ELl eye length in lateral view Ewl eye width in lateral view HLl head length in lateral view HWl head width in lateral view LLMPl length of lateral margin pronotum in lateral view BLl body length including wings in lateral view LAWl length of the anterior wing in lateral view WLHd width-length radio of head in dorsal view WLPd width-length ratio of pronotum in dorsal view WLSd width-lengh radio of scutellum in dorsal view WLCv width-length radio of clypeus PRv postclypeus ratio in ventral view (width/length) SRv stylet ratio in ventral view (width/length) RCRv coxa ratio (width/length) RBWl ratio between body length with wings and length without wings RPCl postclypeus radio in lateral view (width/length) Rel eye ratio in lateral view (width/length) HRALl head ratio in lateral view (width/length) BLWl ratio between the length of body with wings and forewing length in lateral view.
11) Head width with eyes (HWEd) , 12) head length in dorsal view (HLd), 13) pronotum width in dorsal view (PWd), 14) pronotum length in dorsal view (PLd), 15) scutellum width in dorsal view (SWd), 16) scutellum length in dorsal view (SLd), 17) postclypeus length in ventral view (PcLv), 18) postclypeus width in ventral view (PcWv), 19) anteclypeus length in ventral view (Alv), 20) anteclypeus width in ventral view (AWv), 21) stylet length in ventral view (SLv), 22) stylet width in ventral view (SWv), 23) posterior coxa width in ventral view (PCWv), 24) posterior coxa length in ventral view (PcLv), 25) body length without wings in ventral view (BLWv), 26) postclypeus length in lateral view (PCLl), 27) postclypeus width in lateral view (PCWl), 28) eye length in lateral view (ELl), 29) eye width in lateral view (Ewl), 30) head length in lateral view (HLl), 31) head width in lateral view (HWl), 32) length of lateral margin pronotum in lateral view (LLMPl), 33) body length including wings in lateral view (BLl), 34) length of the anterior wing in lateral view (LAWl), 35) width-length radio of head in dorsal view (WLHd), 36) width-length ratio of pronotum in dorsal view (WLPd), 37) width-lengh radio of scutellum in dorsal view (WLSd), 38) width-length radio of clypeus (WLCv), 39) postclypeus ratio in ventral view (width/length) (PRv), 40) stylet ratio in ventral view (width/length) (SRv), 41) coxa ratio (width/length) (RCRv), 42) ratio between body length with wings and length without wings (RBWl), 43) postclypeus radio in lateral view (width/length) (RPCl), 44) eye ratio in lateral view (width/length) (REl), 45) head ratio in lateral view (width/length) (HRALl), 46) and ratio between the length of body with wings and forewing length in lateral view (BLWl).
The frequency of character states for each feature was calculated for each taxon in the contingency tables (Tables
Frequencies of multi-state or binary characters used to compare the variation of color patterns of tegmen among Mexican Aeneolamia species. Abbreviations: TC tegmen color CIE color of internal clavus edge in the tegmen CAT color pattern on the anterior third of tegmen CDT color pattern on the distal third of tegmen.
Attribute | Character states | A. danpecki | A. albofasciata | A. contigua | Chi2: | CC** |
---|---|---|---|---|---|---|
1.- TC | (1) Black. | 50 (78%) | 362 (73%) | 0 | 180.5* | 0.62 |
(2) Dark brown. | 14 (22%) | 134 (27%) | 48 (70%) | |||
(3) Light brown. | 0 | 0 | 20 (30%) | |||
2.- CIE | (1) Same color as tegmen. | 64 (100%) | 397 (80%) | 0 | 300.94* | 0.73 |
(2) With a yellowish or white line. | 0 | 99 (20%) | 0 | |||
(3) With an orange line. | 0 | 0 | 68 (100%) | |||
3.- CAT | (1) The same color as tegmen. | 54 (85%) | 0 | 0 | 500* | 0.81 |
(2) Incomplete thin transversal line. | 10 (15%) | 0 | 0 | |||
(3) Complete broad transversal yellowish or white line. | 0 | 494 (100%) | 0 | |||
(4) Complete broad transversal orange or red line. | 0 | 0 | 68 (100%) | |||
4.- CDT | (1) The same color as tegmen. | 54 (85%) | 0 | 0 | 600* | 0.81 |
(2) With an incomplete thin transversal line. | 10 (15%) | 0 | 0 | |||
(3) With a complete broad transversal. yellowish or white line. | 0 | 494 (100%) | 0 | |||
(4) With a complete broad transversal orange or red line. | 0 | 0 | 68 (100%) | |||
n = | 64 | 494 | 68 |
Frequencies of multi-state or binary characters used to compare the variation male genitalia among Mexican Aeneolamia species. Abbreviations: EAE elevation of the anal tube sclerites SGP shape of subgenital plates SEGP shape of internal edge of subgenital plate apex ASP paramere, shape of primary apical spine SSP parameter, shape of secondary subapical spine PRE tip of aedeagus spines.
Attribute | Character state | A. danpecki | A. albofasciata | A. contigua | Chi2: | CC** |
---|---|---|---|---|---|---|
5.- EAE | (1) Tenth and eleventh tergites at the same level on the horizontal. | 0 | 7 (58%) | 0 | ||
(2) Tenth higher than the eleventh tergite, with respect to the horizontal. | 10 (100%) | 5 (42%) | 7 (100%) | 143.8* | 0.56 | |
6.- SGP | (1) Acute apex with straight lateral edges. | 10 (100%) | 12(100%) | 0 | ||
(2) Apex obliquely truncate with lateral edges slightly concave. | 0 | 0 | 7 (100%) | |||
7.- SEGP | (1) Acuminate in a pointed lobe. | 10 (100%) | 0 | 0 | ||
(2) Acuminate in a rounded lobe. | 0 | 12 (100%) | 0 | |||
(3) Blunt. | 0 | 0 | 7 (100%) | 300* | 0.7 | |
8.- ASP | (1) Long and thin spine with a continuous curvature not angulated. | 10 (100%) | 12 (100%) | 0 | ||
(2) Short and wide spine with a pronounced angulated curvature. | 0 | 0 | 7 (100%) | 300* | 0.7 | |
9.- SSP | (1) With two rounded acute lobes similar in size and shape. | 10 (100%) | 0 | 0 | ||
(2) With two acute lobes, the dorsal one conspicuously bigger than the ventral. | 0 | 12 (100%) | 0 | |||
(3) With one big lobe. | 0 | 0 | 7 (100%) | 600* | 0.81 | |
10.- PRE | (1) Conspicuously curved upward. | 10 (100%) | 0 | 0 | ||
(2) Slightly curved upward. | 0 | 12 (100%) | 0 | |||
(3) Slightly curved downward. | 0 | 0 | 7 (100%) | 414* | 0.81 | |
n = | 10 | 11 | 7 |
Measurements of morphological characteristics of three Mexican Aeneolamia spp. as mean ± standard deviation (mm); * Features that display statistically significant differences among species supported by two way ANOVA; in these cases mean values with the same letter were not significantly different at the 5% level by the Tukey test. Abbreviations: HWEd Head width with eyes HLd head length in dorsal view, PWd pronotum width in dorsal view PLd pronotum length in dorsal view SWd scutellum width in dorsal view SLd scutellum length in dorsal view PcLv postclypeus length in ventral view PCWv postclypeus width in ventral view Alv anteclypeus length in ventral view AWv anteclypeus width in ventral view SLv stylet length in ventral view SWv stylet width in ventral view PcWv posterior coxa width in ventral view PcLv posterior coxa length in ventral view BLWv body length without wings in ventral view PCLl postclypeus length in lateral view PCWl postclypeus width in lateral view ELl eye length in lateral view Ewl eye width in lateral view HLl head length in lateral view HWl head width in lateral view LLMPl length of lateral margin pronotum in lateral view BLl body length including wings in lateral view LAWl length of the anterior wing in lateral view WLHd width-length radio of head in dorsal view WLPd width-length ratio of pronotum in dorsal view WLSd width-lengh radio of scutellum in dorsal view WLCv width-length radio of clypeus PRv postclypeus ratio in ventral view (width/length) SRv stylet ratio in ventral view (width/length) RCRv coxa ratio (width/length) RBWl ratio between body length with wings and length without wings RPCl postclypeus radio in lateral view (width/length) REl eye ratio in lateral view (width/length) HRALl head ratio in lateral view (width/length) BLWl ratio between the length of body with wings and forewing length in lateral view.
Attribute Abbreviation (mm) | A. danpecki | A. albofasciata | A. contigua |
---|---|---|---|
11.-HWEd | 1.90 ± 0.07 | 2.04 ± 0.03 | 1.96 ± 0.07 |
12.-HLd | 0.87 ± 0.03 | 0.94 ± 0.01 | 0.91 ± 0.04 |
13.-PWd | 2.73 ± 0.10 | 2.90 ± 0.04 | 2.81 ± 0.10 |
14.-PLd | 1.62 ± 0.06 | 1.75 ± 0.02 | 1.68 ± 0.06 |
15.-SWd | 1.24 ± 0.05 | 1.18 ± 0.01 | 1.14 ± 0.04 |
16.-SLd* | 1.43c ± 0.05 | 1.57b ± 0.02 | 1.61a ± 0.02 |
17.-PcLV | 1.24 ± 0.04 | 1.30 ± 0.02 | 1.22 ± 0.04 |
18.-PCWv | 0.83 ± 0.05 | 0.83 ± 0.02 | 0.77 ± 0.03 |
19.-Alv* | 0.70b ± 0.02 | 0.76a ± 0.01 | 0.78a ± 0.03 |
20.-AWv | 0.48 ± 0.02 | 0.53 ± 0.02 | 0.48 ± 0.02 |
21.-SLv* | 0.76b ± 0.03 | 0.77b ± 0.01 | 0.82a ± 0.01 |
22.-SWv* | 0.19b ± 0.00 | 0.22a ± 0.01 | 0.21a ± 0.01 |
23.-PcWv | 0.49 ± 0.02 | 0.54 ± 0.01 | 0.52 ± 0.02 |
24.-PcLv | 0.59 ± 0.02 | 0.62 ± 0.01 | 0.59 ± 0.02 |
25.-BLWv* | 6.33c ± 0.23 | 7.01a ± 0.09 | 6.67b ± 0.23 |
26.-PCLI | 0.73 ± 0.03 | 0.76 ± 0.02 | 0.71 ± 0.02 |
27.-PCWI | 1.17 ± 0.04 | 1.24 ± 0.03 | 1.18 ± 0.02 |
28.-ElI | 0.47 ± 0.02 | 0.51 ± 0.01 | 0.49 ± 0.02 |
29.-EwI | 0.63 ± 0.02 | 0.68 ± 0.01 | 0.67 ± 0.02 |
30.-HLI | 0.90 ± 0.03 | 0.92 ± 0.02 | 0.87 ± 0.03 |
31.-HWI | 1.34 ± 0.05 | 1.40 ± 0.02 | 1.38 ± 0.02 |
32.-LLMPI* | 0.65c ± 0.03 | 0.72a ± 0.02 | 0.68b ± 0.02 |
33.-BLI | 7.91 ± 0.27 | 8.21 ± 0.12 | 8.21 ± 0.28 |
34.-LAWI | 6.42 ± 0.22 | 6.57 ± 0.08 | 6.64 ± 0.22 |
35.-WLHd | 0.14 ± 0.00 | 0.14 ± 0.00 | 0.13 ± 0.01 |
36.-WLPd | 0.10 ± 0.00 | 0.11 ± 0.00 | 0.10± 0.00 |
37.-WLSd | 0.05 ± 0.00 | 0.05 ± 0.00 | 0.05 ± 0.00 |
38.-WLCv | 0.09 ± 0.00 | 0.10 ± 0.00 | 0.1 ± 0.00 |
39.-PRv | 0.09 ± 0.00 | 0.10 ± 0.00 | 0.1 ± 0.00 |
40.-SRv | 0.25 ± 0.01 | 0.24 ± 0.00 | 0.25 ± 0.01 |
41.-RCRv | 0.05 ± 0.00 | 0.06 ± 0.00 | 0.05 ± 0.00 |
42.-RBWl | 0.08 ± 0.00 | 0.08 ± 0.00 | 0.07 ± 0.00 |
43.-RPCl | 0.04 ± 0.00 | 0.04 ± 0.00 | 0.03 ± 0.00 |
44.-REl | 0.05 ± 0.00 | 0.05 ± 0.00 | 0.04 ± 0.00 |
45.-HRALl | 0.05 ± 0.00 | 0.04 ± 0.00 | 0.04 ± 0.00 |
46.-BLWl | 0.08 ± 0.00 | 0.08 ± 0.00 | 0.07 ± 0.00 |
Measurements of morphological characteristics of three Mexican Aeneolamia spp. (Mean and standard deviation (mm)); * Features that display significant statistical differences among interaction species-sexes supported by two-way ANOVA, in each species, mean values with the same letter were not significantly different at the 5% level by Tukey test. Abbreviations: HWEd Head width with eyes HLd head length in dorsal view, PWd pronotum width in dorsal view PLd pronotum length in dorsal view SWd scutellum width in dorsal view SLd scutellum length in dorsal view PcLv postclypeus length in ventral view PCWv postclypeus width in ventral view Alv anteclypeus length in ventral view AWv anteclypeus width in ventral view SLv stylet length in ventral view SWv stylet width in ventral view PcWv posterior coxa width in ventral view PcLv posterior coxa length in ventral view BLWv body length without wings in ventral view PCLl postclypeus length in lateral view PCWl postclypeus width in lateral view ELl eye length in lateral view Ewl eye width in lateral view HLl head length in lateral view HWl head width in lateral view LLMPl length of lateral margin pronotum in lateral view BLl body length including wings in lateral view LAWl length of the anterior wing in lateral view WLHd width-length radio of head in dorsal view WLPd width-length ratio of pronotum in dorsal view WLSd width-lengh radio of scutellum in dorsal view WLCv width-length radio of clypeus PRv postclypeus ratio in ventral view (width/length) SRv stylet ratio in ventral view (width/length) RCRv coxa ratio (width/length) RBWl ratio between body length with wings and length without wings RPCl postclypeus radio in lateral view (width/length) REl eye ratio in lateral view (width/length) HRALl head ratio in lateral view (width/length) BLWl ratio between the length of body with wings and forewing length in lateral view.
Attribute Acronym (mm) | A. danpecki | A. albofasciata | A. contigua | |||
---|---|---|---|---|---|---|
♀ | ♂ | ♀ | ♂ | ♀ | ♂ | |
11.-HWEd* | 2.06a ± 0.01 | 1.75b ± 0.12 | 2.13 ± 0.028 | 1.95 ± 0.04 | 2.11a ± 0.02 | 1.81b ± 0.12 |
12.-HLd* | 0.95a ± 0.01 | 0.79b ± 0.05 | 1.00 ± 0.014 | 0.89 ± 0.01 | 1.02a ± 0.05 | 0.81b ± 0.06 |
13.-PWd* | 2.95a ± 0.03 | 2.50b ± 0.17 | 3.01 ± 0.039 | 2.78 ± 0.05 | 3.00 ± 0.04 | 2.62 ± 0.18 |
14.-PLd | 1.74 ± 0.02 | 1.49 ± 0.10 | 1.82 ± 0.028 | 1.68 ± 0.03 | 1.80 ± 0.02 | 1.57 ± 0.11 |
15.-SWd* | 1.35a ± 0.04 | 1.13b ± 0.08 | 1.23 ± 0.015 | 1.13 ± 0.02 | 1.20 ± 0.02 | 1.09 ± 0.08 |
16.-SLd* | 1.58a ± 0.03 | 1.28b ± 0.09 | 1.64 ± 0.031 | 1.49 ± 0.02 | 1.61 ± 0.02 | 1.44 ± 0.11 |
17.-PcLV* | 1.35a ± 0.01 | 1.13b ± 0.08 | 1.37 ± 0.028 | 1.22 ± 0.02 | 1.29 ± 0.03 | 1.15 ± 0.08 |
18.-PCWv* | 0.97a ± 0.06 | 0.69b ± 0.05 | 0.91 ± 0.025 | 0.75 ± 0.02 | 0.85a ± 0.03 | 0.70b ± 0.05 |
19.-Alv* | 0.74 ± 0.01 | 0.66 ± 0.05 | 0.79 ± 0.019 | 0.73 ± 0.02 | 0.86a ± 0.01 | 0.72b ± 0.05 |
20.-AWv* | 0.54a ± 0.01 | 0.42b ± 0.03 | 0.58a ± 0.025 | 0.47b ± 0.01 | 0.55a ± 0.01 | 0.43b ± 0.03 |
21.-SLv* | 0.82a ± 0.02 | 0.71b ± 0.05 | 0.81 ± 0.021 | 0.73 ± 0.01 | 0.84 ± 0.02 | 0.75 ± 0.06 |
22.-SWv | 0.20 ± 0.00 | 0.20 ± 0.00 | 0.23 ± 0.009 | 0.21 ± 0.00 | 0.23 ± 0.01 | 0.19 ± 0.01 |
23.-PcWv* | 0.55a ± 0.01 | 0.43b ± 0.03 | 0.58 ± 0.011 | 0.50 ± 0.01 | 0.57a ± 0.01 | 0.43b ± 0.04 |
24.-PcLv* | 0.64 ± 0.01 | 0.55 ± 0.04 | 0.65 ± 0.013 | 0.59 ± 0.02 | 0.64a ± 0.01 | 0.50b ± 0.05 |
25.-BLWv* | 6.91a ± 0.08 | 5.70b ± 0.39 | 7.27 ± 0.115 | 6.74 ± 0.11 | 7.15 ± 0.10 | 6.21 ± 0.43 |
26.-PCLI* | 0.79a ± 0.02 | 0.68b ± 0.05 | 0.85a ± 0.017 | 0.67b ± 0.00 | 0.78a ± 0.02 | 0.66b ± 0.02 |
27.-PCWI* | 1.27a ± 0.02 | 1.07b ± 0.07 | 1.32a ± 0.033 | 1.16b ± 0.03 | 1.22 ± 0.02 | 1.14 ± 0.03 |
28.-ElI* | 0.50 ± 0.01 | 0.44 ± 0.03 | 0.54 ± 0.008 | 0.48 ± 0.01 | 0.54a ± 0.01 | 0.45b ± 0.03 |
29.-EwI* | 0.68 ± 0.01 | 0.61 ± 0.04 | 0.71a ± 0.010 | 0.65b ± 0.01 | 0.73 ± 0.01 | 0.63 ± 0.04 |
30.-HLI* | 0.98a ± 0.02 | 0.83b ± 0.06 | 0.98 ± 0.019 | 0.86 ± 0.01 | 0.95 ± 0.01 | 0.81 ± 0.06 |
31.-HWI* | 1.47a ± 0.01 | 1.22b ± 0.08 | 1.48 ± 0.033 | 1.33 ± 0.02 | 1.45 ± 0.02 | 1.32 ± 0.02 |
32.-LLMPI* | 0.72a ± 0.02 | 0.58b ± 0.04 | 0.76 ± 0.020 | 0.68 ± 0.02 | 0.73 ± 0.02 | 0.63 ± 0.04 |
33.-BLI* | 8.39a ± 0.07 | 7.45b ± 0.51 | 8.56 ± 0.125 | 7.85 ± 0.16 | 8.66a ± 0.11 | 7.79b ± 0.54 |
34.-LAWI* | 6.81a ± 0.04 | 6.04b ± 0.42 | 6.84 ± 0.075 | 6.31 ± 0.10 | 6.94 ± 0.06 | 6.35 ± 0.44 |
35.-WLHd | 0.14 ± 0.00 | 0.15 ± 0.00 | 0.14 ± 0.002 | 0.15 ± 0.00 | 0.14 ± 0.00 | 0.14 ± 0.01 |
36.-WLPd | 0.11 ± 0.00 | 0.11 ± 0.00 | 0.11 ± 0.001 | 0.11 ± 0.00 | 0.11 ± 0.00 | 0.10 ± 0.01 |
37.-WLSd | 0.06 ± 0.00 | 0.06 ± 0.00 | 0.05 ± 0.003 | 0.05 ± 0.00 | 0.05 ± 0.00 | 0.05 ± 0.00 |
38.-WLCv | 0.10 ± 0.00 | 0.10 ± 0.01 | 0.10 ± 0.002 | 0.11 ± 0.00 | 0.10 ± 0.00 | 0.11 ± 0.01 |
39.-PRv | 0.09 ± 0.00 | 0.10 ± 0.01 | 0.09 ± 0.003 | 0.10 ± 0.00 | 0.11 ± 0.00 | 0.10 ± 0.01 |
40.-SRv | 0.27a ± 0.01 | 0.23b ± 0.02 | 0.24 ± 0.007 | 0.24 ± 0.01 | 0.25 ± 0.01 | 0.25 ± 0.02 |
41.-RCRv* | 0.06a ± 0.00 | 0.05b ± 0.00 | 0.06 ± 0.001 | 0.06 ± 0.00 | 0.06a ± 0.00 | 0.05b ± 0.01 |
42.-RBWl | 0.08 ± 0.00 | 0.08 ± 0.01 | 0.08 ± 0.001 | 0.08 ± 0.00 | 0.08 ± 0.00 | 0.08 ± 0.01 |
43.-RPCl | 0.04 ± 0.00 | 0.04 ± 0.00 | 0.04 ± 0.001 | 0.04 ± 0.00 | 0.04 ± 0.00 | 0.04 ± 0.00 |
44.-REl | 0.05 ± 0.00 | 0.04 ± 0.00 | 0.05 ± 0.001 | 0.05 ± 0.00 | 0.05 ± 0.00 | 0.05 ± 0.00 |
45.-HRALl | 0.05 ± 0.00 | 0.05 ± 0.00 | 0.04 ± 0.001 | 0.04 ± 0.00 | 0.04 ± 0.00 | 0.04 ± 0.00 |
46.-BLWl | 0.08 ± 0.00 | 0.08 ± 0.00 | 0.08 ± 0.001 | 0.08 ± 0.00 | 0.08 ± 0.00 | 0.08 ± 0.01 |
Results from two-way ANOVA to compare the variation of measurements of morphological characteristics of three Mexican Aeneolamia spp. Abbreviations: HWEd Head width with eyes HLd head length in dorsal view, PWd pronotum width in dorsal view PLd pronotum length in dorsal view SWd scutellum width in dorsal view SLd scutellum length in dorsal view PcLv postclypeus length in ventral view PCWv postclypeus width in ventral view Alv anteclypeus length in ventral view AWv anteclypeus width in ventral view SLv stylet length in ventral view SWv stylet width in ventral view PcWv posterior coxa width in ventral view PcLv posterior coxa length in ventral view BLWv body length without wings in ventral view PCLl postclypeus length in lateral view PCWl postclypeus width in lateral view ELl eye length in lateral view Ewl eye width in lateral view HLl head length in lateral view HWl head width in lateral view LLMPl length of lateral margin pronotum in lateral view BLl body length including wings in lateral view LAWl length of the anterior wing in lateral view WLHd width-length radio of head in dorsal view WLPd width-length ratio of pronotum in dorsal view WLSd width-lengh radio of scutellum in dorsal view WLCv width-length radio of clypeus PRv postclypeus ratio in ventral view (width/length) SRv stylet ratio in ventral view (width/length) RCRv coxa ratio (width/length) RBWl ratio between body length with wings and length without wings RPCl postclypeus radio in lateral view (width/length) REl eye ratio in lateral view (width/length) HRALl head ratio in lateral view (width/length) BLWl ratio between the length of body with wings and forewing length in lateral view.
Acronym | Species | Sex | Interaction | |||
---|---|---|---|---|---|---|
F | P | F | P | F | P | |
11.-HWEd | 1.744 | 0.181 | 18.75 | 4.09E-05 | 0.5288 | 0.5913 |
12.-HLd | 1.69 | 0.191 | 24.43 | 3.89E-06 | 0.7839 | 0.4599 |
13.-PWd | 1.261 | 0.289 | 16.7 | 9.95E-05 | 0.532 | 0.5894 |
14.-PLd | 2.074 | 0.132 | 15.43 | 0.0001752 | 0.4199 | 0.6585 |
15.-SWd | 2.127 | 0.126 | 12.87 | 0.0005607 | 0.9456 | 0.3925 |
16.-SLd | 7.015 | 0.002 | 24.03 | 4.64E-06 | 3.41 | 0.03773 |
17.-PcLV | 1.247 | 0.293 | 17.76 | 6.27E-05 | 0.3953 | 0.6747 |
18.-PCWv | 1.284 | 0.282 | 37.3 | 3.03E-08 | 1.751 | 0.1799 |
19.-Alv | 4.34 | 0.016 | 14.45 | 0.000272 | 0.8141 | 0.4465 |
20.-AWv | 2.79 | 0.067 | 41.35 | 7.38E-09 | 0.07588 | 0.927 |
21.-SLv | 3.333 | 0.041 | 14.03 | 0.0003309 | 0.9593 | 0.3874 |
22.-SWv | 5.43 | 0.006 | 10.72 | 0.001547 | 0.362 | 0.6974 |
23.-PcWv | 2.785 | 0.068 | 35.9 | 5.47E-08 | 0.5938 | 0.5546 |
24.-PcLv | 0.8573 | 0.428 | 16.48 | 0.0001129 | 0.5701 | 0.5677 |
25.-BLWv | 3.915 | 0.024 | 19.2 | 3.38E-05 | 0.923 | 0.4013 |
26.-PCLI | 1.625 | 0.203 | 40.45 | 1.01E-08 | 1.033 | 0.3603 |
27.-PCWI | 2.054 | 0.135 | 23.38 | 5.94E-06 | 1.596 | 0.2088 |
28.-ElI | 2.106 | 0.128 | 18.46 | 4.65E-05 | 0.4292 | 0.6524 |
29.-EwI | 1.037 | 0.359 | 12.17 | 0.0007772 | 0.2618 | 0.7703 |
30.-HLI | 0.7906 | 0.457 | 22.52 | 8.46E-06 | 0.06886 | 0.9335 |
31.-HWI | 1.068 | 0.348 | 30.43 | 3.75E-07 | 1.291 | 0.2803 |
32.-LLMPI | 3.131 | 0.049 | 22.86 | 7.33E-06 | 0.6679 | 0.5155 |
33.-BLI | 0.5747 | 0.565 | 10.47 | 0.001739 | 0.06293 | 0.939 |
34.-LAWI | 0.3988 | 0.672 | 9.184 | 0.003245 | 0.1178 | 0.889 |
35.-WLHd | 0.2057 | 0.815 | 0.1278 | 0.7216 | 0.4334 | 0.6498 |
36.-WLPd | 0.2475 | 0.781 | 2.128 | 0.1484 | 0.543 | 0.583 |
37.-WLSd | 1.834 | 0.166 | 1.272 | 0.2626 | 0.02747 | 0.9729 |
38.-WLCv | 0.7389 | 0.481 | 2.875 | 0.09368 | 0.02094 | 0.9793 |
39.-PRv | 2.144 | 0.124 | 2.638 | 0.1081 | 1.154 | 0.3202 |
40.-SRv | 2.506 | 0.088 | 1.278 | 0.2615 | 4.683 | 0.01183 |
41.-RCRv | 2.852 | 0.063 | 9.638 | 0.002617 | 0.9715 | 0.3828 |
42.-RBWl | 1.17 | 0.315 | 0.551 | 0.46 | 0.3265 | 0.7223 |
43.-RPCl | 0.4304 | 0.652 | 2.741 | 0.1015 | 0.1586 | 0.8536 |
44.-REl | 2.267 | 0.11 | 2.545 | 0.1144 | 1.71 | 0.1871 |
45.-HRALl | 0.2143 | 0.808 | 0.381 | 0.5388 | 0.4524 | 0.6377 |
46.-BLWl | 0.8206 | 0.444 | 2.027 | 0.1582 | 0.6383 | 0.5307 |
To explore if the variation of morphological characteristics together segregates the specimens of A. danpecki sp. nov. in a discrete group within multidimensional spaces, a series of ordination analyses were performed. A principal coordinate analysis (PCoA) was performed from a Gower pairwise matrix among 28 male specimens using the ten discrete (male tegmina color pattern and male genitalia) and 36 continuous features. Also, three principal components analyses (PCAs) were performed to explore the geographical patterns of morphological variation among specimens using pairwise covariance matrices of 36 continuous characters. Additionally, we include canonical variate analyses (CVAs) to determine to what extent these features explained the possible taxonomic segregation based on the 90 specimens in males, females, and both sexes together. Multivariate analyses were performed considering each specimen as an operational taxonomic unit (OTU). Lastly, we looked for multivariate statistical differences among taxonomic groups of Aeneolamia recovered in the ordination analyses, with an analysis of similarities (ANOSIM) and the respective pairwise Hotelling’s T non-parametric tests among groups representing putative species. Groups recovered in the multivariate space were confirmed by the comparative morphological analysis of male genitalia.
From the male genitalia images that show the aedeagus intact, shape variation in patterns of aedeagus spines were quantified among A. danpecki sp. nov. (n = 4), A. albofasciata (n = 6), and A. contigua (n = 7) specimens using potential homologous landmarks (lm) and semi-landmarks (sml) (
To illustrate the geographic distribution of Aeneolamia spp., the records of the analyzed specimens were projected onto a map of Mexican biogeographical provinces (
In total, 46 morphological characters were evaluated: four discrete characters focused on tegmen color patterns in both sexes, six discrete characters on male genitalia, and 36 continuous characters were measured in specimens of both sexes: Six continuous quantitative morphological were reported by
All tegmina and male genitalia features showed differences in character state frequencies among A. danpecki sp. nov., A. albofasciata, and A. contigua (Tables
Aeneolamia danpecki: dark brown (22%) to black (78%) tegmen (Fig.
Aeneolamia albofasciata: dark brown (27%) to black (73%) tegmen (Fig.
Aeneolamia contigua: light brown (30%) to dark brown (70%) tegmen (Fig.
Combining morphometric data of both sexes, Aeneolamia danpecki is smaller than both A. albofasciata and A. contigua in most features analyzed except SWd, and HRALl (Table
Two-way ANOVA also supported significant statistical differences between sexes in more than 20 features (Tables
The first two principal coordinates of PCoAs of continuous and discrete features of males explained 65% of variations (PCo1 = 39.99%, PCo2 = 15.01%) (Fig.
Scatter plots of ordination analyses to evaluate the morphological variation of Mexican Aeneolamia spp.: A principal coordinate analysis using the 10 discrete (male tegmina color pattern and male genitalia) and 36 continuous features of males B analysis of canonical variation with 36 morphological continuous features of both sexes C females D males. In the center of the scatter plot, vectors corresponding to the contribution of the traits in the multivariate space (B, C, D). The largest circles are the centroid of the polygons (B). Abbreviations: Alv Anteclypeus length in ventral view BLs body length without wings in ventral view BLWv body length without wings in ventral view LAWl length of the anterior wing in lateral view PCWv postclypeus width in ventral view PLd postclypeus length in dorsal view PWd pronotum width in dorsal view PcWs postclypeus width in lateral view SWd scutellum width in dorsal view SLv stylet length in ventral view.
The superimposition of 15 aedeagus spine configurations of Aeneolamia members (A. albofasciata, n = 5; A. contigua, n = 6; A. danpecki, n = 4) showed that shape variation is found on both proximal and medial regions (Fig.
Scatter plots among the first three relative warps with its respective deformation grids ± 1.5 SD, corresponding to shape analysis of aedeagus spine (sp) of Mexican Aeneolamia spp. A Rw1 vs. Rw2 B Rw1 vs. Rw3 C position of landmarks (S1 and S2) and semi-landmarks (1–14) on aedeagus spine of A. contigua D deformation grids ± 1.5 SD.
Because our analyses support qualitative and quantitative discrete phenotypic variation among Aeneolamia species (two tegmina features characters and five genitalia ones) and the most pronounced morphological differences compared to the previously recognized species A. albofasciata and A. contigua, the specimens of the new species are grouped into a new taxon, Aeneolamia danpecki Castro, Armendáriz, Utrera, sp. nov., described below.
Holotype. HOM-TIP-166, 1 ♂ adult, coll. U. Castro-Valderrama and Youssef Utrera-Vélez leg., 28 September 2003, on Paspalum sp., Sola de Vega, 16°27'44.48"N, 97°1'25.73"W, 1715 m a.s.l., Oaxaca state, Mexico. Pinned adult deposited in
Paratypes. HOM-TIP-167, 1 ♀, same data as holotype; 1 ♀, coll. Cervantes, A. Delgado, C. Mayorga, S. Gámez leg.; 5 km W San Martín Lachila, Mpio Zimatlán, Oaxaca, México, 16°35'39.18"N, 96°52'14.16"W, 12 July 2004. Pinned specimens deposited in
The epithet is a noun in the nominative singular standing in apposition to the genus Aeneolamia, in honor of Dr. Daniel C. Peck for his contributions to the knowledge of Cercopidae and his friendship with UC-V.
Aeneolamia danpecki Castro, Armendáriz, Utrera, sp. nov. is assigned to the genus Aeneolamia by virtue of its tubular aedeagus with a single pair of slender spines attached anteriorly near the middle of the shaft. It can be distinguished from the other known Mexican species of Aeneolamia by the following combination of characters: tegmen dark brown to black, with two incomplete and barely visible transverse bands, one oblique band on the basal third, and another straight band on the distal third or only basal band visible or both absent (Figs
Male adult of A. danpecki (Holotype). Sola de Vega, Oaxaca A dorsal view B ventral view C lateral view D head in dorsal view E head in ventral view F prothorax in dorsal view G head in lateral view H abdomen in ventral view I anterior section of wing J mounted holotype K genital vial and labels of holotype.
Female adult of A. danpecki (Paratype). Sola de Vega, Oaxaca A dorsal view B ventral view C lateral view D head in dorsal view E head in ventral view F prothorax in dorsal view G anterior section of wing H abdomen in ventral view I head in lateral view J median section of wing K distal section of wing.
Male measurements. Lateral view length (N = 15) 7.45 ± 0.51 mm; width of head in dorsal view (N = 15) 1.75 ± 0.12 mm.
Head. Dorsal view (Fig.
Thorax. Dorsal view (Fig.
Abdomen. Ventral view (Fig.
Genitalia. Pygofer in lateral view (Figs
Female measurements. Lateral view length (N = 15) 8.39 ± 0.07 mm; width of head in dorsal view (N = 15) 2.06 ± 0.01 mm. Same characteristics as the male, except larger and posterior and lateral edges of each sternite light brown or reddish (Fig.
Paspalum sp. and Pennisetum sp.
Aeneolamia danpecki has black or dark brown subgenital plates with an acute end. In the type of material, San Martín Lachila is a municipality and not part of the Municipality of Zimatlán. Aeneolamia danpecki was recognized as distinct for the first time as “Aeneolamia aff. albofasciata (Lallemand, 1939)” by
1 | Apex of subgenital plates obliquely truncate (Fig. |
2 |
– | Apex of the subgenital plates acute (Fig. |
3 |
2 | Tegmen light brown to dark brown, with two orange transverse lines (Fig. |
A. contigua (Walker, 1851) |
– | Tegmen black with narrow oblique transverse basal line interrupted at claval suture, a distal line straight, with lines on claval edges V-shaped (see |
A. contigua campecheana Fennah, 1951 |
3 | Tegmen dark brown to black, with one or two yellowish or white transverse lines (Fig. |
A. albofasciata (Lallemand, 1939) |
– | Tegmen dark brown to black, with two incomplete and barely visible transverse bands, one oblique band on basal third and another straight band on the distal third or only basal band visible or both absent (Figs |
A. danpecki Castro, Armendáriz & Utrera, sp. nov. |
The distribution of A. danpecki was supported by three occurrence records from Sierra Madre del Sur province, in Oaxaca state (Fig.
The evaluation of ten discrete characters of male tegmen and genitalia indicates that six of them (CAT, CDT, SGP, SEGP, SSP, and PRE) are useful to differentiate A. danpecki, and both sets of features together can differentiate this species from A. albofasciata and A. contigua as well as being diagnostic characters for A. danpecki (Tables
The statistical analysis of morphological variation of Mexican members of Aeneolamia supports the earlier suggestion that specimens identified previously as A. aff. albofasciata in
From quantitative continuous and discrete characters (PCoA) of males and quantitative features of both sexes (PCA), permitted the recovery of discrete groups corresponding to the two previously recognized species, A. albofasciata and A. contigua, and the new species A. danpecki (Figs
Morphometric analyses have been poorly explored in Cercopidae and quantitative analyses of the shape of genital structures have not been performed previously in the family. However, in studies of other Cicadomorpha and other Cercopoidea, morphometric analyses have been used to recognize and delimit new species. In the genus Cycloscytina Martynov, 1926, shape analysis of the wing allowed to elucidate the species status of its members and support an extinct new species from the Triassic (
In this study, it is evident that, at the biogeographical level, A. danpecki sp. nov. is in sympatry with A. contigua and A. albofasciata in the Sierra Madre del Sur. However, the records of Mexican Aeneolamia species (Table
This research was supported by Programa para el Desarrollo Profesional Docente, Tipo Superior-2019 (PRODEP), project “Taxonomía y distribution de la familia Cercopidae (Hemiptera: Auchenorrhyncha) en México”, granted to UC-V, and PAPIIT-UNAM IA201720, IA203122 and CONACyT Fronteras de la Ciencia (139030) granted to FA-T. Likewise, Grupo Papalotla S. A. de C. V. for supporting the trip to Oaxaca where A. danpecki was collected in 2003. We thank Cristina Mayorga Martínez (