Research Article |
Corresponding author: Santiago Niño-Maldonado ( coliopteranino@hotmail.com ) Academic editor: Astrid Eben
© 2016 Uriel Jeshua Sánchez-Reyes, Santiago Niño-Maldonado, Ludivina Barrientos-Lozano, Shawn M. Clark, Robert W. Jones.
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:
Sánchez-Reyes UJ, Niño-Maldonado S, Barrientos-Lozano L, Clark SM, Jones RW (2016) Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational and temporal gradients in Sierra de San Carlos, Mexico. ZooKeys 611: 11-56. https://doi.org/10.3897/zookeys.611.9608
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The study of biodiversity of Chrysomelidae in Mexico and its variation within ecological gradients has increased recently, although important areas in the country remain to be explored. We conducted a faunistic inventory and analyzed the elevational and temporal variation of leaf beetle communities in the Sierra de San Carlos, in the state of Tamaulipas, in northeastern Mexico. This is an area with high to extreme priority for conservation, and due to its insular geographical position and to the vegetational communities present, it must be considered as a sky island. We selected seven sample sites distributed in different elevations within three localities, and comprising different vegetational communities. At each site, we randomly delimited 12 sample plots of 400 m2 where sampling was conducted by entomological sweep netting and collecting directly by hand. Sampling was conducted monthly at each plot, for a total of 1,008 samples between February 2013 and January 2014. By the end of the study, we had obtained a total of 3,081 specimens belonging to six subfamilies, 65 genera, and 113 species, with Trichaltica scabricula (Crotch, 1873) being recorded for first time in Mexico. Species richness was less than the values observed at other studies conducted in the same region, which is attributed to differences in the number of plant species and to the insular location of Sierra de San Carlos; however, the higher diversity values suggest a higher quality of natural resources and vegetational communities. No consistent pattern of leaf beetle communities was correlated with elevation, although higher values of species richness and diversity were obtained at the highest elevation site. The seasonal gradient showed that the rainy season is most favorable for leaf beetle communities. We found that species composition was different between sites and months, and also that there exists a significant association between the abundance obtained at each site and particular months. These results highlight the importance of different microhabitats for species distribution, and suggest that each species of Chrysomelidae has a differential response to environmental factors that vary within the elevational gradient and according to seasons. Also, we confirm and emphasize the important status of Sierra de San Carlos as a key natural area for biological conservation.
Biodiversity, chrysomelid beetles, ecological gradient, elevation, seasonality, sky island
Chrysomelidae (excluding Bruchinae or seed beetles), whose members are also known as leaf beetles, is one of the most diverse taxa within Coleoptera, with more than 35,000 to 40,000 described species worldwide (
Mexico is located within an important geographic area, and the inhabiting fauna is the result of the interface of the Neotropical and Nearctic realms. So, the study of chrysomelid distribution in this region is useful to analyze the biogeographical and ecological patterns of its species in the American continent. In Mexico, the most explored and studied areas are the Baja California peninsula (
Recently, a series of faunistic and ecological studies on leaf beetle fauna has been conducted in the northeastern portion of Mexico, specifically in the state of Tamaulipas. To date, 250 species have been recorded from this state, which now ranks fourth in chrysomelid diversity from Mexico (
Although data from faunistic inventories constitute a very important descriptor of diversity and allow analysis of species distribution from a region, it is also important that the variation of ecological patterns are associated with natural gradients, as they reflect the ecological and evolutionary adaptations of species to various environmental conditions (
The study was conducted in the Sierra de San Carlos (Figure
One of the most important characteristics of Sierra de San Carlos is its designation as a Priority Conservation Area in Mexico due to its biological, ecological and physiogeographical features: it is the northern limit of the Cloud Forest vegetation in Mexico, and it has some endemic plant species; also, it is considered as a biogeographical island (“sky island”) due to its isolation from other nearby mountain ranges, such as the Sierra Madre Oriental and Sierra de Tamaulipas (
We selected seven sampling sites distributed in three localities within Sierra de San Carlos (Figure
Twelve sampling plots of 400 m2 each (20 × 20 m) were established within each of the seven sampling sites. Plot dimensions were determined with the species-area curve method, using the nested quadrat type (
Sampling data in the Cerro El Diente locality, Sierra de San Carlos, Mexico (coordinates at center of plot; elevation in meters).
Cerro El Diente | |||||||
---|---|---|---|---|---|---|---|
Site 1 – Submountain scrub | Site 2 – Tamaulipan thorny scrub | ||||||
Sampling plot | Latitude | Longitude | Elevation | Sampling plot | Latitude | Longitude | Elevation |
P1 | 24°33.020'N | 98°57.004'W | 550 | P1 | 24°32.468'N | 98°57.454'W | 772 |
P2 | 24°33.048'N | 98°57.013'W | 548 | P2 | 24°32.471'N | 98°57.402'W | 790 |
P3 | 24°33.062'N | 98°56.960'W | 544 | P3 | 24°32.471'N | 98°57.374'W | 784 |
P4 | 24°33.068'N | 98°57.047'W | 547 | P4 | 24°32.492'N | 98°57.353'W | 766 |
P5 | 24°33.104'N | 98°57.073'W | 535 | P5 | 24°32.501'N | 98°57.383'W | 773 |
P6 | 24°32.996'N | 98°57.096'W | 555 | P6 | 24°32.490'N | 98°57.416'W | 778 |
P7 | 24°32.936'N | 98°57.082'W | 561 | P7 | 24°32.496'N | 98°57.460'W | 760 |
P8 | 24°32.920'N | 98°57.173'W | 570 | P8 | 24°32.522'N | 98°57.469'W | 750 |
P9 | 24°33.031'N | 98°57.333'W | 571 | P9 | 24°32.537'N | 98°57.473'W | 743 |
P10 | 24°33.105'N | 98°57.316'W | 557 | P10 | 24°32.531'N | 98°57.452'W | 750 |
P11 | 24°32.995'N | 98°57.237'W | 560 | P11 | 24°32.522'N | 98°57.423'W | 755 |
P12 | 24°33.046'N | 98°57.137'W | 540 | P12 | 24°32.543'N | 98°57.441'W | 745 |
Site 3 – Oak forest | Site 4 – Cloud forest | ||||||
Sampling plot | Latitude | Longitude | Elevation | Sampling plot | Latitude | Longitude | Elevation |
P1 | 24°32.038'N | 98°57.496'W | 938 | P1 | 24°31.780'N | 98°57.557'W | 1077 |
P2 | 24°32.018'N | 98°57.466'W | 935 | P2 | 24°31.795'N | 98°57.565'W | 1065 |
P3 | 24°32.021'N | 98°57.500'W | 950 | P3 | 24°31.780'N | 98°57.593'W | 1070 |
P4 | 24°32.026'N | 98°57.539'W | 948 | P4 | 24°31.774'N | 98°57.622'W | 1055 |
P5 | 24°31.996'N | 98°57.489'W | 964 | P5 | 24°31.753'N | 98°57.631'W | 1085 |
P6 | 24°31.984'N | 98°57.456'W | 948 | P6 | 24°31.760'N | 98°57.672'W | 1077 |
P7 | 24°31.973'N | 98°57.480'W | 971 | P7 | 24°31.744'N | 98°57.695'W | 1093 |
P8 | 24°32.002'N | 98°57.543'W | 967 | P8 | 24°31.730'N | 98°57.738'W | 1112 |
P9 | 24°31.995'N | 98°57.516'W | 974 | P9 | 24°31.738'N | 98°57.783'W | 1109 |
P10 | 24°31.968'N | 98°57.527'W | 993 | P10 | 24°31.751'N | 98°57.816'W | 1102 |
P11 | 24°31.981'N | 98°57.550'W | 982 | P11 | 24°31.776'N | 98°57.831'W | 1086 |
P12 | 24°31.970'N | 98°57.577'W | 979 | P12 | 24°31.793'N | 98°57.868'W | 1076 |
Sampling data in the Ejido Carricitos y Tinajas locality, Sierra de San Carlos, Mexico (coordinates at center of plot; elevation in meters).
Ejido Carricitos y Tinajas | |||||||
---|---|---|---|---|---|---|---|
Site 5 – Riparian and secondary vegetation | Site 6 – Oak and pine forests | ||||||
Sampling plot | Latitude | Longitude | Elevation | Sampling plot | Latitude | Longitude | Elevation |
P1 | 24°35.807'N | 99°2.450'W | 700 | P1 | 24°35.397'N | 99°3.037'W | 839 |
P2 | 24°35.789'N | 99°2.484'W | 701 | P2 | 24°35.420'N | 99°3.023'W | 830 |
P3 | 24°35.764'N | 99°2.508'W | 704 | P3 | 24°35.440'N | 99°3.041'W | 816 |
P4 | 24°35.727'N | 99°2.534'W | 712 | P4 | 24°35.463'N | 99°3.017'W | 814 |
P5 | 24°35.684'N | 99°2.600'W | 716 | P5 | 24°35.491'N | 99°3.028'W | 795 |
P6 | 24°35.719'N | 99°2.654'W | 720 | P6 | 24°35.567'N | 99°3.067'W | 813 |
P7 | 24°35.673'N | 99°2.766'W | 740 | P7 | 24°35.563'N | 99°3.101'W | 827 |
P8 | 24°35.632'N | 99°2.851'W | 757 | P8 | 24°35.575'N | 99°3.124'W | 846 |
P9 | 24°35.605'N | 99°2.894'W | 764 | P9 | 24°35.579'N | 99°3.146'W | 860 |
P10 | 24°35.571'N | 99°2.870'W | 763 | P10 | 24°35.577'N | 99°3.175'W | 866 |
P11 | 24°35.545'N | 99°2.806'W | 773 | P11 | 24°35.548'N | 99°3.013'W | 788 |
P12 | 24°35.533'N | 99°2.909'W | 776 | P12 | 24°35.584'N | 99°2.970'W | 780 |
Sampling data in the San Nicolás locality, Sierra de San Carlos, Mexico (coordinates at center of plot; elevation in meters).
Site 7 – Tamaulipan thorny scrub and submountain scrub vegetation | |||
---|---|---|---|
Sampling plot | Latitude | Longitude | Elevation |
P1 | 24°32.356'N | 98°46.936'W | 502 |
P2 | 24°32.319'N | 98°47.006'W | 501 |
P3 | 24°32.290'N | 98°47.073'W | 500 |
P4 | 24°32.224'N | 98°47.117'W | 499 |
P5 | 24°32.180'N | 98°47.153'W | 499 |
P6 | 24°32.371'N | 98°46.883'W | 503 |
P7 | 24°32.391'N | 98°46.840'W | 508 |
P8 | 24°32.427'N | 98°46.797'W | 510 |
P9 | 24°32.450'N | 98°46.759'W | 508 |
P10 | 24°32.485'N | 98°46.702'W | 508 |
P11 | 24°32.334'N | 98°46.922'W | 502 |
P12 | 24°32.295'N | 98°46.906'W | 503 |
Systematic sampling was conducted between 10:00 and 17:00 h, using a standard entomological sweep net of 40 cm diameter. Individual samples consisted of 120-200 sweeps of the shrub and herbaceous vegetation in each plot. Contents of the net were emptied into a plastic bag, adding 60% ethanol and an indelible label with corresponding data. Each plot (12) within the seven sites was sampled monthly, from February 2013 to January 2014, comprising 1,008 total samples at the end of the study; sweeping was conducted by the same person during the whole study to reduce sampling error. Each sample and the specimens obtained were processed according to the method described by
Identification of specimens was made using available literature on Chrysomelidae (
We obtained environmental data from two meteorological stations located in the municipalities of San Carlos and San Nicolás in the study area. Historical data of total monthly rainfall and monthly average temperature (only the average from 1951-2010 was available) were plotted to visually analyze the fluctuation of these parameters. On this basis, four seasons were defined: Early dry season (EDS: November, December, January), Late dry season (LDS: February, March, April), Early rainy season (ERS: May, June, July), and Late rainy season (LRS: August, September, October). Data of precipitation and temperature were correlated with species richness and abundance, using a Spearman correlation analysis in STATISTICA 8.0 (
All the following analyses were made only with the data obtained through systematic sampling (i.e., by sweeping in 12 plots at each site). Species collected otherwise were excluded from the analysis, but are included in the checklist of species.
As a measure of species richness, we used the total number of species present throughout the Sierra de San Carlos, and at each site and season. Significant differences in the number of species were assessed through permutation tests in PAST 3.07 (
Species from Sierra de San Carlos were divided into five categories, according to their total abundance: 1) very common (more than 70 individuals); 2) common (11 to 70 specimens); 3) rare (3 to 10 specimens); 4) doubletons (two specimens); and 5) singletons (one specimen only). These categories were used because they have been implemented in similar studies with Chrysomelidae, also in the state of Tamaulipas (
Finally, the association of abundance and species richness obtained at each elevational site during each month was measured with a Correspondence analysis. This is a multivariate technique based on contingency tables and count data, where the significant statistical dependence between rows (sites) and columns (months) is tested by a chi-square test (
In total, 3,081 specimens, belonging to 109 species, 63 genera and six subfamilies, were obtained. An additional four species were obtained by independent collecting in various sites from study area, resulting in a total species richness of 113 species and 65 genera within the study period (2013-2014) in the Sierra de San Carlos (Appendix
Examples of leaf beetle biodiversity from Sierra de San Carlos, Mexico. A Lema balteata LeConte, 1884 B Lema opulenta Harold, 1874 C Helocassis clavata (Fabricius, 1798) D Plagiodera semivittata Stål, 1860 E Miraces aeneipennis Jacoby, 1888 F Malacorhinus acaciae (Schaeffer, 1906)G Cyclotrypema furcata (Olivier, 1808) H Acrocyum dorsalis Jacoby, 1885 I Colaspis melancholica Jacoby, 1881 J Griburius montezuma (Suffrian, 1852) K Cryptocephalus trizonatus Suffrian, 1858 L Coscinoptera tamaulipasi Medvedev, 2012 M Diplacaspis prosternalis (Schaeffer, 1906).
Examples of leaf beetle biodiversity from Sierra de San Carlos, Mexico. A Trichaltica scabricula (Crotch, 1873), new country record. In decreasing order from B to I, the most abundant species in the current study. B Syphrea sp. 2 C Diachus sp. 1 D Xanthonia sp. 1 E Centralaphthona diversa (Baly, 1877) F Chrysogramma sp. 1 G Pachybrachis sp. 1 H Sumitrosis inaequalis (Weber, 1801) I Margaridisa atriventris (Melsheimer, 1847).
Observed and estimated species richness of Chrysomelidae by site and season at Sierra de San Carlos, Mexico. Elevation in meters. EDS = Early dry season, LDS = Late dry season, ERS = Early rainy season, LRS = Late rainy season.
Sobs | Nonparametric indexes | Clench model | Completeness (%) | ||||
---|---|---|---|---|---|---|---|
Chao 1 | Jack 1 | ACE | Sest | Slope | |||
Elevation* | |||||||
500 (S7) | 33 d | 36.5±3.5 | 42.93±3.35 | 38.7±0.65 | 42.07 | 0.05 | 76.8–90.4 |
550 (S1) | 38 abcd | 41.57±3.1 | 48.92±3.75 | 47.58±0 | 53.82 | 0.07 | 77.6–91.4 |
730 (S5) | 36 defg | 52.9±12.72 | 49.9±4.06 | 50.57±0.71 | 48.31 | 0.066 | 68.05–72.1 |
760 (S2) | 40 ae | 40.89±1.26 | 46.95±2.57 | 43.43±0.46 | 50.51 | 0.05 | 85.19–97.82 |
820 (S6) | 27 cg | 39.25±13.15 | 33.95±2.93 | 33.62±0.71 | 31.47 | 0.03 | 68.78–79.52 |
960 (S3) | 47 bf | 50.38±3.06 | 56.93±3.35 | 52.83±0.53 | 60.83 | 0.074 | 82.55–93.29 |
1080 (S4) | 50 b | 58.64±6.82 | 62.91±5.44 | 58.76±0.49 | 60.89 | 0.065 | 79.47–85.26 |
Season* | |||||||
EDS | 40 a | 44±3.74 | 48.96±2.94 | 45.88±0.5 | 47.5 | 0.026 | 81.69–90.9 |
LDS | 49 b | 55.4±5.92 | 57.96±3.26 | 53.55±0.27 | 57.49 | 0.029 | 84.54–88.44 |
ERS | 78 c | 91.88±8.32 | 102.9±5.69 | 94.66±1.17 | 102.17 | 0.075 | 75.8–84.89 |
LRS | 76 c | 82.62±4.57 | 93.93±4.54 | 87.65±0.77 | 92.97 | 0.058 | 80.91–91.98 |
Total | 109 | 115.88±4.5 | 128.98±5.06 | 122.47±0 | 120.46 | 0.012 | 84.5–94 |
The most abundant subfamily in the study area was Galerucinae (including Alticini), with 53.6% (1,652 specimens) of the total abundance of Chrysomelidae in the Sierra de San Carlos, followed by Cryptocephalinae with 26.9% (828 specimens). Less abundance was found in Eumolpinae (12.6%, 388 specimens), Cassidinae (4.9%, 152 specimens), Chrysomelinae (1.1%, 33 specimens) and Criocerinae (0.9%, 28 specimens). Species counts per subfamily were greatest for Galerucinae, with 54 species (49.5%) representing half of the total species richness recorded in Sierra de San Carlos. Lower values were recorded for Cryptocephalinae with 27 species (24.8%), Eumolpinae with nine species (8.3%), Cassidinae and Criocerinae both with eight species (7.3%), and Chrysomelinae with only three species (2.8%).
Eight species were categorized as “very abundant species,” each with over 70 specimens that accounted for 60.3% (1,859 total specimens) of the total abundance obtained from Sierra de San Carlos. Of these eight species, Syphrea sp. 2 (475 specimens) and Diachus sp. 1 (418) were the most abundant, followed by Xanthonia sp. 1 (276), Centralaphthona diversa (Baly, 1877) (244), Chrysogramma sp. 1 (193), Pachybrachis sp. 1 (103), Sumitrosis inaequalis (Weber, 1801) (78), and Margaridisa atriventris(Melsheimer, 1847) (72) (Figure
No clear patterns of species richness were found with elevation. The greatest number of species (50) was recorded at the site of highest elevation (1080 masl), but this value was not significantly different from values observed at 960 (47 species) and 550 masl (38 species). The smallest number of species (27) was registered at a high elevation site (820 masl); however, it was not significantly different from sites at 730 (36 species) and 550 masl (Table
Regarding seasonal analysis, the species richness increased progressively and significantly from early dry season (40 species) to early rainy season (78 species). The value decreased to 76 species during late rainy season, although this change was not significant. Seasonal values of estimated species richness through nonparametric indexes and the Clench model followed the same pattern as that of sites, because all values were above 70% of completeness, with slopes under 0.1 for all the seasons (Table
We found significant variations in abundance and diversity of Chrysomelidae between sites of differing elevation (Kruskal Wallis Test, H=100.7, p<0.0001). However, these parameters did not show a specific trend with the increase or decrease in elevation. For example, the highest abundance (665 individuals) was present at the lowest site (500 masl), whereas the lowest value (173 individuals) was obtained at the second lowest elevation (550 masl). Also, differences in abundance obtained between the site of lowest elevation (665 individuals) and site at 960 masl (561 individuals) were not significant, while the number of specimens (440) at the highest site was not statistically different from values observed at lower sites (960, 820 and 760 masl). The lowest abundances were present at 550 (173 specimens) and 730 masl (231 specimens), and these values were significantly different from other elevational sites (Tables
Mann-Whitney pairwise comparisons of chrysomelid abundance between elevational sites in Sierra de San Carlos, Mexico. Upper diagonal = Mann-Whitney U values. Lower diagonal = p values; marked values (*) are significant.
S1–550 m | S2–760 m | S3–960 m | S4–1080 m | S5–730 m | S6–820 m | S7–500 m | |
---|---|---|---|---|---|---|---|
Site 1 | – | 6723 | 5344 | 6289 | 8234 | 6197 | 4675 |
Site 2 | <0.0001* | – | 8853 | 9920 | 8336 | 9735 | 8126 |
Site 3 | <0.0001* | 0.03007* | – | 9232 | 6782 | 9509 | 9634 |
Site 4 | <0.0001* | 0.52 | 0.1042 | – | 7778 | 10100 | 8530 |
Site 5 | 0.001504* | 0.003257* | <0.0001* | 0.000181* | – | 7707 | 6002 |
Site 6 | <0.0001* | 0.3634 | 0.2197 | 0.748 | 0.000121* | – | 8758 |
Site 7 | <0.0001* | 0.001351* | 0.2958 | 0.008679* | <0.0001* | 0.02159* | – |
As observed with abundance, no clear patterns of diversity were found with elevation, although all sites were significantly different from each other. The highest dominance (D=0.406) and lowest entropy (H’=1.716) were obtained in the site of lowest elevation (500 m), indicating that the lowest diversity was found at this site (1/D=2.46; eH’=5.56). However, the second highest value of diversity in the study area (1/D=12.66; and eH’=21.47) was obtained at the second lowest elevation site (550 meters). Diversity decreased progressively from the third (730 meters) to the fifth elevational site (820 m), and then increased from 960 masl to the highest elevation site (1080 masl), where the lowest dominance (D=0.0508) and highest values of entropy (H’=3.328) and diversity (1/D=19.66; eH’=27.88) were obtained (Table
Elevational variation of abundance and diversity of Chrysomelidae in Sierra de San Carlos, Mexico.
Parameter | Study site ‡ | ||||||
---|---|---|---|---|---|---|---|
S7 | S1 | S5 | S2 | S6 | S3 | S4 | |
Elevation (masl) | 500 | 550 | 730 | 760 | 820 | 960 | 1080 |
Abundance † | 665 a | 173 b | 231 c | 432 d | 579 df | 561 aef | 440 de |
Diversity * | |||||||
D = Simpson index (dominance) | 0.406 a | 0.078 b | 0.101 c | 0.137 d | 0.183 e | 0.156 f | 0.0508 g |
1/D = Simpson Diversity index | 2.46 a | 12.66 b | 9.89 c | 7.29 d | 5.45 e | 6.38 f | 19.66 g |
H´ = Shannon index | 1.716 a | 3.067 b | 2.819 c | 2.638 d | 2.185 e | 2.576 d | 3.328 g |
eH´ = Shannon Diversity | 5.56 a | 21.47 b | 16.76 c | 13.98 d | 8.89 e | 13.14 d | 27.88 g |
According to the PERMANOVA analysis, the leaf beetle composition between sites was statistically different (SStotal=33.16; SSwithin-group=23.22; F=5.492, p=0.0001), and almost all pairwise comparisons were significantly different, except for Site 2 (760 masl) and Site 3 (960 masl) (F=1.595, p=0.1151) (Table
PERMANOVA pairwise comparisons of chrysomelid composition between elevational sites in Sierra de San Carlos, Mexico. Upper diagonal = F values. Lower diagonal = p values.
S7–500 m | S1–550 m | S5–730 m | S2–760 m | S6–820 m | S3–960 m | S4–1080 m | |
---|---|---|---|---|---|---|---|
Site 7 | – | 7.446 | 11.65 | 10.89 | 12.17 | 9.959 | 9.959 |
Site 1 | 0.0001 | – | 4.59 | 3.452 | 5.391 | 4.057 | 4.692 |
Site 5 | 0.0001 | 0.0001 | – | 5.13 | 2.497 | 4.229 | 4.658 |
Site 2 | 0.0001 | 0.0001 | 0.0001 | – | 4.171 | 1.595 | 3.449 |
Site 6 | 0.0001 | 0.0001 | 0.0182 | 0.0001 | – | 3.231 | 3.636 |
Site 3 | 0.0001 | 0.0001 | 0.0001 | 0.1151 | 0.0002 | – | 2.644 |
Site 4 | 0.0001 | 0.0001 | 0.0001 | 0.0001 | 0.0001 | 0.0007 | – |
Bray–Curtis similarity between elevational sites in Sierra de San Carlos, Mexico. Upper diagonal = Index values. Lower diagonal = values expressed as percentage of similarity.
S1–550 m | S2–760 m | S3–960 m | S4–1080 m | S5–730 m | S6–820 m | S7–500 m | |
---|---|---|---|---|---|---|---|
Site 1 | – | 0.25455 | 0.14441 | 0.16639 | 0.24752 | 0.16755 | 0.16468 |
Site 2 | 25.455 | – | 0.59819 | 0.31651 | 0.19005 | 0.36993 | 0.074749 |
Site 3 | 14.441 | 59.819 | – | 0.37163 | 0.18434 | 0.45965 | 0.03752 |
Site 4 | 16.639 | 31.651 | 37.163 | – | 0.27422 | 0.33562 | 0.050679 |
Site 5 | 24.752 | 19.005 | 18.434 | 27.422 | – | 0.45432 | 0.051339 |
Site 6 | 16.755 | 36.993 | 45.965 | 33.562 | 45.432 | – | 0.067524 |
Site 7 | 16.468 | 7.4749 | 3.752 | 5.0679 | 5.1339 | 6.7524 | – |
Abundance and diversity values showed significant variation between seasons (H=92.29, p<0.0001). The lowest number of specimens (433) was recorded at early dry season, and abundance increased significantly at the end of the season (888 specimens). Abundance decreased at early rainy season (690 specimens), although this reduction was not significant; then, the value increased significantly at the late rainy season, which recorded the highest abundance (1,070 specimens) in this study (Tables
Mann-Whitney pairwise comparisons for chrysomelid abundance between seasons in Sierra de San Carlos, Mexico. Upper diagonal = Mann-Whitney U values. Lower diagonal = p values; marked values (*) are significant.
Early dry season | Late dry season | Early rainy season | Late rainy season | |
---|---|---|---|---|
Early dry season | – | 22600 | 23100 | 16400 |
Late dry season | <0.0001* | – | 30900 | 25400 |
Early rainy season | <0.0001* | 0.597 | – | 24300 |
Late rainy season | <0.0001* | 0.000083* | 0.0000038* | – |
Seasonal variation of abundance and diversity of Chrysomelidae in Sierra de San Carlos, Mexico.
Parameter | Dry Season | Rainy Season | ||
---|---|---|---|---|
Early (EDS) | Late (LDS) | Early (ERS) | Late (LRS) | |
Abundance ‡ | 433 a | 888 b | 690 b | 1070 c |
Diversity* | ||||
D = Simpson index (dominance) | 0.146 a | 0.101 b | 0.053 c | 0.104 b |
1/D = Simpson Diversity index | 6.84 a | 9.90 b | 18.68 c | 9.59 b |
H´ = Shannon index | 2.736 a | 2.834 b | 3.508 c | 3.091 d |
eH´ = Shannon Diversity | 15.42 a | 17.01 b | 33.38 c | 21.99 d |
Spearman correlation analysis for abundance and species richness of Chrysomelidae with temperature and precipitation in Sierra de San Carlos, Mexico. Marked values (*) are significant.
Temperature [°C] | Precipitation [mm] | |||
---|---|---|---|---|
San Carlos | San Nicolás | San Carlos | San Nicolás | |
Abundance | 0.532 | 0.559 | 0.531 | 0.349 |
Species Richness | 0.809* | 0.822* | 0.826* | 0.710* |
Seasonal species composition and faunistic similarity were analyzed using monthly values. Significant differences in species composition between months were found (SStotal=33.16; SSwithin-group=27.33; F=1.395, p=0.0009). Pairwise comparisons showed that differences were obtained principally between early (January, February, March, April and May) and later months (July, September, October, November and December) (Table
PERMANOVA pairwise comparisons of chrysomelid composition between months in Sierra de San Carlos, Mexico. Upper diagonal = F values. Lower diagonal = p values; values in bold are significant.
Jan | Feb | Mar | Apr | May | Jun | Jul | Aug | Sep | Oct | Nov | Dec | |
Jan | – | 0.964 | 0.344 | 0.063 | 0.105 | 0.312 | 0.029 | 0.076 | 0.017 | 0.022 | 0.010 | 0.031 |
Feb | 0.191 | – | 0.799 | 0.055 | 0.093 | 0.299 | 0.040 | 0.162 | 0.036 | 0.029 | 0.014 | 0.039 |
Mar | 1.076 | 0.629 | – | 0.666 | 0.350 | 0.115 | 0.031 | 0.179 | 0.023 | 0.014 | 0.010 | 0.033 |
Apr | 1.668 | 1.583 | 0.812 | – | 0.939 | 0.206 | 0.038 | 0.075 | 0.009 | 0.004 | 0.012 | 0.047 |
May | 1.442 | 1.407 | 1.076 | 0.493 | – | 0.587 | 0.287 | 0.163 | 0.008 | 0.006 | 0.014 | 0.035 |
Jun | 1.147 | 1.168 | 1.416 | 1.23 | 0.902 | – | 0.467 | 0.685 | 0.083 | 0.082 | 0.005 | 0.013 |
Jul | 1.749 | 1.607 | 1.639 | 1.563 | 1.12 | 0.995 | – | 0.879 | 0.546 | 0.479 | 0.071 | 0.078 |
Aug | 1.566 | 1.33 | 1.34 | 1.495 | 1.264 | 0.844 | 0.638 | – | 0.739 | 0.627 | 0.012 | 0.032 |
Sep | 1.89 | 1.713 | 1.819 | 1.973 | 1.786 | 1.459 | 0.933 | 0.709 | – | 0.962 | 0.107 | 0.088 |
Oct | 1.89 | 1.8 | 1.988 | 2.08 | 1.831 | 1.486 | 0.986 | 0.821 | 0.041 | – | 0.078 | 0.101 |
Nov | 1.916 | 1.809 | 1.722 | 1.718 | 1.778 | 1.929 | 1.491 | 1.864 | 1.544 | 1.624 | – | 0.972 |
Dec | 1.678 | 1.607 | 1.562 | 1.448 | 1.552 | 1.725 | 1.461 | 1.672 | 1.502 | 1.528 | 0.106 | – |
Bray-Curtis similarity between months in Sierra de San Carlos, Mexico. Upper diagonal = Index values. Lower diagonal = values expressed as percentage of similarity.
Jan | Feb | Mar | Apr | May | Jun | Jul | Aug | Sep | Oct | Nov | Dec | |
Jan | - | 0.721 | 0.487 | 0.36 | 0.398 | 0.433 | 0.298 | 0.356 | 0.295 | 0.309 | 0.230 | 0.273 |
Feb | 72.1 | - | 0.636 | 0.347 | 0.35 | 0.367 | 0.313 | 0.390 | 0.309 | 0.318 | 0.203 | 0.215 |
Mar | 48.7 | 63.6 | - | 0.563 | 0.482 | 0.340 | 0.319 | 0.401 | 0.353 | 0.365 | 0.161 | 0.176 |
Apr | 36.0 | 34.7 | 56.3 | - | 0.690 | 0.403 | 0.325 | 0.294 | 0.264 | 0.291 | 0.140 | 0.176 |
May | 39.8 | 35 | 48.2 | 69.0 | - | 0.509 | 0.314 | 0.267 | 0.248 | 0.262 | 0.150 | 0.144 |
Jun | 43.3 | 36.7 | 34.0 | 40.3 | 50.9 | - | 0.367 | 0.382 | 0.331 | 0.349 | 0.161 | 0.186 |
Jul | 29.8 | 31.3 | 31.9 | 32.5 | 31.4 | 36.7 | - | 0.584 | 0.567 | 0.580 | 0.368 | 0.353 |
Aug | 35.6 | 39.0 | 40.1 | 29.4 | 26.7 | 38.2 | 58.4 | - | 0.573 | 0.547 | 0.259 | 0.277 |
Sep | 29.8 | 30.9 | 35.3 | 26.4 | 24.8 | 33.1 | 56.7 | 57.3 | - | 0.894 | 0.446 | 0.368 |
Oct | 30.9 | 31.8 | 36.5 | 29.1 | 26.2 | 34.9 | 58.0 | 54.7 | 89.4 | - | 0.422 | 0.428 |
Nov | 23.0 | 20.3 | 16.1 | 14.0 | 15.0 | 16.1 | 36.8 | 25.9 | 44.6 | 42.2 | - | 0.698 |
Dec | 27.3 | 21.5 | 17.6 | 17.6 | 14.4 | 18.6 | 35.3 | 27.7 | 36.8 | 42.8 | 69.8 | - |
The Correspondence analysis showed no significant associations in the number of species obtained by site for each month (Total Inertia=0.08796, Chi²=74.147, df=66, p=0.23014). However, the association of abundance between sites and months was significant (Total Inertia=0.32237, Chi²=993.24, df=66, p=0.0000). The most clear associations were observed between abundance obtained at lower elevations (500 and 550 masl) and the period comprised by September to December, while the number of specimens in Site 3 (960 masl) were highly related to August. January and February were principally associated with Site 5 (730 masl) and Site 2 (760 masl). The March-May period was associated with Site 6 (820 masl), and the abundance found at the highest elevation site (1080 masl) was predominantly related with June (Figure
The 113 species of Chrysomelidae recorded in this study document that the Sierra de San Carlos represents a proportion close to 50% of the total leaf beetle species richness presently reported from Tamaulipas (
Although the sampled area for both studies was almost the same (33,600 m2 in Sierra de San Carlos vs. 37,500 m2 in Peregrina Canyon), the total and site-season inventory completeness in Peregrina Canyon was close to 70% (
Elevation is one of the most important factors driving ecological communities, because the abiotic factors and biotic variables together modify species richness and composition of assemblages. Recent evidence suggests that the most common elevational pattern is the increase of diversity and species richness at intermediate elevations (
Although a consistent elevational pattern was not found, the high proportion of inventory completeness through all methods employed indicates that the faunistic composition obtained at each site is representative; so, the values of abundance and diversity were reliable (
When analyzing species composition and beta diversity between sites, we observed that Sites 2 and 3 were the only sites with the same composition and a high faunistic similarity, while almost all other comparisons were different, which is contrary to other findings that show a high similarity of Lepidoptera between elevation and seasons at Cerro El Diente (
Regarding seasonal analysis, patterns observed at Sierra de San Carlos were different from those recorded in the most related study in Peregrina Canyon, Tamaulipas (
According to cluster analysis, three groups were formed, based on faunistic similarity between months: November and December, January to June, and July to October. This inconsistency between the four climate seasons (dry/rainy) and the clustering of months by species compositions is possibly due to differential species responses to seasonal variations, as their temporal niche requirements are very distinct (
In addition to the influence of the geographical location of Sierra de San Carlos, the responses of the chrysomelid community in this study, being elevational, seasonal or both, are suggested to be driven by host plants and vegetational associated variables (
Besides, the results obtained through Correspondence analysis confirm the same tendency of an interaction between abiotic and biotic factors on distribution of Chrysomelidae, because they show that abundance of the leaf beetle community at each site is associated with specific months. This was observed, for example, with the significant association of the lowest sites and abundance obtained at the rainy months, or that observed between Site 3 (960 masl) and August. Consequently, these associations suggest unique and specific temporal-site conditions for the communities of Chrysomelidae, which surely are the result of monthly changes in both environmental (abiotic) conditions and plant variables along the elevational gradient. Since elevational and temporal responses of biological communities arise from the effects (direct or indirect) of these gradients on each species (
The species richness of Chrysomelidae in Sierra de San Carlos was not as high as expected for an area with extreme priority for conservation, which could be the result of the geographical position of the study area. However, the high quality of the vegetational communities is presumably associated to the high diversity values. This is true for the highest site of our elevational gradient, where the highest values of species richness and diversity were obtained, and which must surely be associated with the environmental conditions of the cloud forest vegetation at that site, thus emphasizing the conservation urgency of this relict area and supporting the presence of a sky island within Sierra de San Carlos; hence, its high importance for biological conservation and for investigations of leaf beetle distribution.
The first record of a species for Mexico in Sierra de San Carlos is remarkable. Moreover, many of the specimens here determined as morphospecies could be later recognized as new distribution records, or new species. So, it is possible that leaf beetle species at Sierra de San Carlos constitute a very distinctive faunistic assemblage from other chrysomelid faunas in Mexico, which, added to the absence of a clear elevational pattern, suggests a strong effect of the insular geographical position and other geomorphic characteristics of Sierra de San Carlos on the Chrysomelidae distribution. Rainy season was associated with higher values of the ecological parameters of Chrysomelidae, being consistent with general patterns of temporal distribution of leaf beetles.
Regarding leaf beetle composition, we found evidence that different microhabitats, regardless of the distance, as well as different months, support distinct faunistic assemblages. Most importantly, communities within these particular sites are differentially influenced by changing conditions during seasonal/month variation, as suggested by the Correspondence analysis, and by the direct correlation of temperature and precipitation with species richness. These differences and variations in faunistic composition within the elevational and temporal gradients surely mirror differences in floristic composition and abiotic variables, since both are related to leaf beetle distribution. However, these changes must be addressed at a specific level, because the niche requirements of each species are very distinct. Since this is one of few studies conducted in Mexico concerning chrysomelid biodiversity and the variation along natural gradients, it is important that future research accounts for the specific influence of environmental modification (biotic and abiotic) on chrysomelid species at Sierra de San Carlos and other ecological gradients within Mexico. Also, forthcoming studies must address biogeographical relationships of chrysomelid species existent within this and others areas in the country.
We are grateful to our work crew of the 2012-2014 period, which efficiently assisted in the sampling work: Edmar Meléndez-Jaramillo, Nabil Yessenia Martínez-Ruíz and Brenda Villanueva-Alanís. Also, we thank Vannia del Carmen Gómez-Moreno, Geovany de Jesús Fernández-Azuara, and Luís Castillo, for their general support during field trips to Sierra de San Carlos. Crystian Sadiel Venegas-Barrera provided helpful advice and logistic support for the sampling design, during the preliminary and planning phases of this study, at Instituto Tecnológico de Ciudad Victoria.
The active authorities in 2013 from San Carlos and San Nicolás municipalities granted us permission for fieldwork in different areas within Sierra de San Carlos. We are indebted to Jesús Gutiérrez and Lauro Meléndez de la Serna, who allowed us the access to the Cerro El Diente locality, and also to Ma. del Refugio de la Serna González, Jhanelle Varela de la Serna and Marina Meléndez Vela, for supplying kind support and lodging to the first author, during preliminary fieldwork phases of this project. The first author is grateful to the Consejo Nacional de Ciencia y Tecnología (CONACYT), for a scholarship award granted for M.S. studies at the Instituto Tecnológico de Ciudad Victoria. The authors also thank PROMEP, for additional financial support.
Taxonomic checklist and abundance of Chrysomelidae by site and month in Sierra de San Carlos, Mexico. Site column: numbers in square brackets refer to the plot number where the species was collected within that site; see Material and methods (Tables
Taxon | Site [plot] | Month (abundance) |
---|---|---|
CRIOCERINAE Latreille, 1807 | ||
Tribe Lemini Heinze, 1962 | ||
Lema balteata LeConte, 1884 | Site 4 [4] | Aug (1) |
Lema opulenta Harold, 1874 | Site 5 [6] | Aug (1) |
Lema sp. 1 | Site 3 [2] | Nov (1), Dec (1) |
Neolema sp. 1 | Site 1 [9] | Aug (1) |
Site 2 [4] | Sep (1), Oct (1) | |
Site 3 [2] | Jul (1), Aug (1) | |
Site 4 [4, 7, 8, 9, 10] | Jun (1), Jul (1), Aug (5), Sep (1), Oct (1) | |
Neolema sp. 2 | Site 5 [1] | May (1) |
Oulema sp. 1 | Site 3 [3, 11] | Jul (1), Sep (1), Oct (1) |
Site 5 [3] | Aug (1) | |
Oulema sp. 2 | Site 3 [1, 12] | Aug (1), Sep (1), Oct (1) |
Oulema sp. 3 | Site 4 [1, 7] | Jul (1), Aug (1) |
CASSIDINAE Gyllenhal, 1813 | ||
Tribe Chalepini Weise, 1910 | ||
Brachycoryna pumila Guérin-Méneville, 1844 | Site 1 [1, 2, 4, 7, 9, 11] | May (2), Jun (1), Jul (1), Sep (3), Oct (2) |
Site 2 [1, 2, 3, 6, 7, 9, 11] | Jan (2), Feb (2), Apr (2), Jun (1), Jul (3), Aug (1), Sep (2), Oct (2) | |
Site 4 [3, 4, 9] | May (1), Jul (1), Sep (1), Oct (1), Nov (1), Dec (1) | |
Site 6 [7, 8, 11, 12] | Jan (1), Feb (2), May (1), Jun (2), Jul (1), Aug (4) | |
Site 7 [1, 5] | Jul (1), Aug (1) | |
Chalepus verticalis (Chapuis, 1877) | Site 7 [6] | Sep (1), Oct (1) |
Sumitrosis inaequalis (Weber, 1801) | Site 1 [6, 7, 8] | Jan (1), Feb (1), Jul (1), Sep (1), Oct (1), Nov (2) |
Site 2 [5, 8, 9, 10, 11, 12] | Jan (1), Feb (3), Mar (2), May (1), Jun (2), Jul (3), Aug (1), Sep (2), Oct (1), Nov (6), Dec (2) | |
Site 3 [3, 6, 8] | Jun (1), Jul (1), Sep (1), Oct (1) | |
Site 4 [5, 6, 7, 8, 9, 10, 11, 12] | Jan (1), Feb (1), May (6), Jun (2), Jul (8), Aug (4), Sep (7), Oct (5) | |
Site 5 [2, 8, 11] | Jul (1), Aug (1), Sep (1), Oct (1) | |
Site 6 [1, 3, 9] | May (1), Jul (4) | |
Tribe Mesomphaliini Hope, 1840 | ||
Ogdoecosta juvenca (Boheman, 1854) | Site 4 [8, 9] | Jun (3), Jul (2), Aug (1) |
Tribe Ischyrosonychini Chapuis, 1875 | ||
Physonota alutacea Boheman, 1854 | Site 1 [4] | Sep (1), Oct (1) |
Site 7 [1, 2, 12] | Jun (1), Sep (7), Oct (7), Nov (3), Dec (2) | |
Tribe Cassidini Gyllenhal, 1813 | ||
Coptocycla (Psalidonota) texana (Schaeffer, 1933)* | Site 1 | May |
Helocassis clavata (Fabricius, 1798) | Site 2 [5] | May (1), Jul (1) |
Site 3 [4] | May (1) | |
Site 5 [3] | Jun (1) | |
Helocassis crucipennis (Boheman, 1855) | Site 5 [7] | Aug (1) |
Site 6 [1] | Jul (1) | |
Metrionella bilimeki Spaeth, 1932 | Site 4 [8, 9, 10, 11] | May (1), Jul (3), Aug (3), Sep (4), Oct (3) |
CHRYSOMELINAE Latreille, 1802 | ||
Tribe Chrysomelini Latreille, 1802 | ||
Subtribe Doryphorina Motschulsky, 1860 | ||
Labidomera suturella Chevrolat, 1844 | Site 3 [3] | Nov (1), Dec (1) |
Site 4 [11] | Jul (1) | |
Subtribe Chrysomelina Latreille, 1802 | ||
Plagiodera semivittata Stål, 1860 | Site 2 [5, 10, 12] | Jan (1), Feb (1), Jun (2), Sep (2), Oct (2), Nov (1), Dec (1) |
Site 3 [3, 5, 7, 8] | Jan (1), Feb (1), May (2), Jun (6) | |
Site 4 [2, 3] | Apr (1), Sep (1), Oct (1) | |
Plagiodera thymaloides Stål, 1860 | Site 2 [8, 10, 11] | Jun (2), Jul (1), Aug (2) |
Site 3 [3, 8] | Jun (2) | |
GALERUCINAE Latreille, 1802 | ||
Tribe Galerucini Latreille, 1802 | ||
Group Coelomerites Chapuis, 1875 | ||
Coraia subcyanescens (Schaeffer, 1906) | Site 1 [1] | Jun (1) |
Miraces aeneipennis Jacoby, 1888 | Site 1 [4, 10, 11] | Aug (1), Sep (2), Oct (2), Nov (2), Dec (2) |
Site 2 [3, 4, 7] | Sep (6), Oct (4) | |
Site 3 [4, 9, 10, 12] | Jun (1), Jul (1), Sep (5), Oct (5) | |
Site 7 [12] | Jul (1), Nov (1), Dec (1) | |
Group Schematizites Chapuis, 1875 | ||
Monoxia sp. 1 | Site 1 [4] | Sep (1), Oct (1) |
Ophraella sp. 1 | Site 1 [9] | Jan (1), Feb (1) |
Tribe Metacyclini Chapuis, 1875 | ||
Malacorhinus acaciae (Schaeffer, 1906) | Site 7 [1, 6, 10] | Jun (3), Sep (3), Oct (3) |
Malacorhinus sp. 1 | Site 4 [4] | Nov (1), Dec (1) |
Tribe Luperini Chapuis, 1875 | ||
Subtribe Diabroticina Chapuis, 1875 | ||
Group Diabroticites Chapuis, 1875 | ||
Acalymma invenustum Munroe & Smith, 1980 | Site 1 [5] | Sep (1), Oct (1) |
Site 3 [5] | Sep (1), Oct (1) | |
Site 4 [4] | Aug (2) | |
Gynandrobrotica lepida (Say, 1835) | Site 5 [1, 2, 6, 7, 8, 10, 11, 12] | Apr (1), Aug (3), Sep (11), Oct (9), Nov (6), Dec (5) |
Site 6 [1, 3, 4, 6, 10, 11, 12] | Jul (1), Aug (2), Sep (12), Oct (9), Nov (3), Dec (2) | |
Group Cerotomites Chapuis, 1875 | ||
Cyclotrypema furcata (Olivier, 1808) | Site 4 [6, 10, 12] | Jun (2), Aug (2) |
Site 5 [6] | Aug (1) | |
Tribe Alticini Newman, 1835 | ||
Acallepitrix sp. 1 | Site 1 [12] | Sep (1), Oct (1) |
Site 2 [3, 8] | Jan (2), Feb (2), Nov (1) | |
Site 3 [9] | Mar (1) | |
Site 4 [7, 8] | Mar (1), Aug (1) | |
Site 5 [1, 2, 3, 6, 12] | Jan (1), Feb (1), Mar (1), Aug (3), Sep (5), Oct (3), Nov (2), Dec (2) | |
Site 6 [1, 2, 4, 5, 6, 8, 9] | Jan (3), Feb (8), Mar (11), Apr (1), Jul (2), Aug (2) | |
Acallepitrix sp. 2 | Site 5 [3, 7] | Aug (2) |
Site 6 [1, 2] | Mar (3), Apr (1), May (1), Jul (1) | |
Acallepitrix sp. 3 | Site 4 [4] | Jun (1) |
Acallepitrix sp. 4 | Site 1 [2] | Sep (1), Oct (1) |
Site 3 [2, 3, 11] | Jul (2), Aug (1), Sep (2), Oct (2) | |
Site 4 [2, 4, 6, 7, 8, 9, 11, 12] | Jun (1), Jul (3), Aug (4), Sep (4), Oct (2) | |
Site 5 [3] | Sep (1), Oct (1) | |
Site 6 [8] | Jun (2), Sep (1), Oct (1) | |
Acrocyum dorsalis Jacoby, 1885 | Site 2 [3, 10] | Mar (1), Sep (1), Oct (1) |
Site 3 [2] | Jan (1), Feb (1) | |
Alagoasa decemguttata (Fabricius, 1801) | Site 4 [6, 8] | Apr (1), Sep (1), Oct (1) |
Altica sp. 1 | Site 4 [10] | Jul (1) |
Altica sp. 2 | Site 3 [1, 10] | Jul (1), Aug (1) |
Altica sp. 3 | Site 5 [11] | Apr (1) |
Blepharida rhois (Forster, 1771) | Site 7 [1, 4, 11, 12] | Aug (1), Sep (2), Oct (2), Nov (3), Dec (2) |
Centralaphthona diversa (Baly, 1877) | Site 1 [2, 3, 6, 12] | Jan (2), Feb (3), Jun (2), Aug (1), Sep (1), Oct (1) |
Site 2 [2, 4, 10, 12] | Jan (2), Feb (2), Mar (1), Jun (7), Aug (6), Sep (5), Oct (4), Nov (2), Dec (2) | |
Site 3 [1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12] | Jan (16), Feb (29), Mar (4), May (4), Jul (4), Aug (15), Sep (3), Oct (2) | |
Site 4 [1, 2, 3, 4, 6, 8, 9, 10, 11, 12] | Jan (4), Feb (8), Mar (19), Apr (4), May (2), Jun (4), Jul (3), Aug (5), Sep (2), Oct (2) | |
Site 5 [1, 2, 3, 4, 6, 11] | Jan (2), Feb (2), Apr (1), May (1), Jun (3), Jul (2), Aug (3), Sep (1), Oct (1) | |
Site 6 [1, 4, 5, 6, 7, 8, 9, 10, 11, 12] | Jan (8), Feb (17), Mar (3), Apr (1), May (2), Jun (10), Jul (1), Aug (9), Sep (2), Oct (2) | |
Site 7 [1, 3] | Apr (1), Aug (1) | |
Centralaphthona sp. 2 | Site 2 [3, 4, 11] | Jan (1), Feb (1), Jul (1), Aug (1) |
Site 3 [9] | May (1) | |
Site 6 [9] | May (1) | |
Centralaphthona sp. 3 | Site 3 [3, 7] | Jan (1), Feb (1), Aug (1) |
Site 4 [9, 10] | Jan (1), Feb (1), Mar (1) | |
Site 5 [9, 11, 12] | Jun (1), Aug (2) | |
Site 6 [3, 4] | May (1), Jul (1) | |
Centralaphthona sp. 4 | Site 6 [11] | Jul (1) |
Chaetocnema sp. 1 | Site 1 [2, 3, 7] | Jun (1), Jul (18), Aug (1), Sep (10), Oct (8) |
Site 5 [2, 3, 4] | Aug (3), Sep (3), Oct (3) | |
Site 6 [6, 10, 11] | Aug (1), Sep (4), Oct (3) | |
Chaetocnema sp. 2 | Site 1 [11] | Nov (1), Dec (1) |
Site 2 [10] | Nov (1), Dec (1) | |
Site 3 [3, 8] | Apr (1), Nov (1), Dec (1) | |
Site 4 [5, 10, 11, 12] | Jan (2), Feb (3), Mar (1), Aug (7) | |
Site 5 [2, 11] | Jan (1), Feb (1), Mar (1) | |
Site 6 [6, 9] | Jan (1), Feb (1), Apr (2) | |
Chaetocnema sp. 3 | Site 1 [6] | Sep (1), Oct (1) |
Site 2 [3] | Jan (1), Feb (1) | |
Site 3 [9] | May (1) | |
Site 4 [1, 12] | May (3) | |
Site 5 [2, 4, 5, 6] | Jan (1), Feb (1), Apr (1), May (5), Sep (1), Oct (1) | |
Site 6 [6, 11] | Jan (2), Feb (2), Jun (1) | |
Site 7 [1, 3, 6] | Nov (3), Dec (3) | |
Chaetocnema sp. 4 | Site 6 [11] | May (1) |
Chrysogramma sp. 1 | Site 1 [6] | Apr (1) |
Site 2 [1, 2, 3, 4, 6, 7, 9, 10, 11, 12] | Jan (12), Feb (41), Mar (45), Apr (6), May (2), Jul (1) | |
Site 3 [1, 2, 3, 4, 5, 6, 7, 8, 10, 11, 12] | Jan (14), Feb (34), Mar (26), Apr (8), May (3) | |
Dibolia sp. 1 | Site 4 [4] | Aug (1) |
Disonycha glabrata (Fabricius, 1781) | Site 5 [1, 6] | Aug (2) |
Disonycha stenosticha Schaeffer, 1931 | Site 1 [6] | Nov (2) |
Site 2 [3, 4, 6, 8, 11, 12] | Jul (2), Sep (2), Oct (2), Nov (5), Dec (1) | |
Site 3 [1, 7, 9] | Jul (1), Nov (3), Dec (2) | |
Site 6 [8] | Aug (1) | |
Disonycha teapensis Blake, 1933 | Site 3 [1] | Sep (1), Oct (1) |
Dysphenges sp. 1 | Site 1 [4, 10] | Jul (2) |
Site 3 [1, 5, 8, 9] | Jul (4), Sep (1), Oct (1) | |
Site 7 [2, 9] | Jul (1), Sep (1) | |
Epitrix sp. 1 | Site 3 [8] | Jan (1), Feb (1) |
Site 4 [3, 5, 6, 8, 9, 10] | Jan (1), Feb (1), Mar (2), May (3), Jun (2), Aug (1) | |
Site 5 [1, 2, 3, 4, 6, 7, 8, 12] | Jan (2), Feb (2), Mar (1), May (1), Jun (2), Jul (1), Aug (2), Sep (1), Oct (1) | |
Site 6 [6, 9, 11, 12] | Jan (2), Feb (2), May (1), Aug (1), Sep (3), Oct (1) | |
Epitrix sp. 2 | Site 1 [6] | Nov (1) |
Site 2 [2, 10] | Jan (1), Feb (1), Mar (1), Nov (1), Dec (1) | |
Site 3 [1, 5, 8, 10, 11, 12] | Jan (3), Feb (3), Mar (4), Jul (1), Aug (1) | |
Site 4 [5, 7, 11, 12] | Jan (1), Feb (1), Mar (2), Apr (1) | |
Site 5 [6, 8] | Jan (1), Feb (1), Jun (1) | |
Epitrix sp. 3 | Site 2 [9] | Mar (1) |
Site 4 [1, 9, 12] | Jan (1), Feb (1), Mar (2) | |
Site 7 [6] | Sep (1), Oct (1) | |
Epitrix sp. 4 | Site 5 [3] | Jun (2), Aug (2) |
Hypolampsis sp. 1 | Site 4 [6] | Jun (1) |
Kuschelina laeta (Perbosc, 1839) | Site 7 [6] | Aug (1) |
Longitarsus sp. 1 | Site 1 [6] | Mar (1) |
Site 2 [2] | Jul (2) | |
Site 4 [1, 3] | Mar (1), Apr (1) | |
Site 7 [1, 6] | Jan (1), Feb (1), Sep (1), Oct (1) | |
Longitarsus sp. 2 | Site 1 [8] | May (1) |
Site 2 [2, 5] | Jun (1), Aug (1) | |
Site 3 [3, 4, 5, 8, 10] | Jul (1), Aug (1), Nov (3), Dec (2) | |
Site 4 [1, 3, 4, 5, 7, 8, 9, 10] | Jan (3), Feb (4), Mar (5), Apr (1), May (1), Jun (1), Jul (5), Aug (2), Sep (3), Oct (1) | |
Site 5 [1, 3, 5, 8] | May (1), Aug (3) | |
Site 6 [5, 10] | Jul (2), Aug (1) | |
Site 7 [1] | Jul (1) | |
Lupraea sp. 1 | Site 3 [5] | Sep (1), Oct (1) |
Site 4 [1, 4, 5, 7. 8, 10, 11] | Apr (3), Jul (3), Aug (7), Sep (2), Oct (1) | |
Lupraea sp. 2 | Site 3 [4] | Sep (1), Oct (1) |
Site 4 [1, 4] | Sep (4), Oct (2) | |
Lysathia sp. 1 | Site 5 [6] | Mar (1) |
Margaridisa atriventris (Melsheimer, 1847) | Site 2 [2, 5, 12] | Aug (3) |
Site 3 [1, 4, 5, 6, 8, 10, 11, 12] | Jan (1), Feb (1), May (1), Jun (1), Jul (7), Aug (14), Sep (2), Oct (2) | |
Site 4 [8, 9] | Mar (1), Jul (2) | |
Site 5 [1, 3, 10, 11] | Mar (1), Jun (1), Jul (2), Aug (2), Sep (1), Oct (1) | |
Site 6 [1, 2, 4, 5, 7, 12] | Jan (1), Feb (1), May (1), Jun (2), Jul (18), Aug (5) | |
Site 7 [6] | Aug (1) | |
Monomacra sp. 1 | Site 4 [7] | May (2) |
Parchicola sp. 1 | Site 2 [2, 5, 7] | Jul (3), Aug (1), Sep (1), Oct (1) |
Site 3 [1, 10, 12] | Jul (4) | |
Phydanis nigriventris Jacoby, 1891 | Site 3 [3] | Aug (1) |
Site 7 [11] | Aug (1) | |
Phyllotreta sp. 1 | Site 1 [5, 9] | Mar (2) |
Site 2 [7] | Jan (1), Feb (1) | |
Site 3 [2, 11, 12] | Mar (3), May (1) | |
Site 4 [7, 9] | Mar (4), May (1) | |
Site 5 [1] | Apr (1) | |
Site 6 [6, 9, 10] | Mar (15) | |
Plectrotetra sp. 1 | Site 3 [3] | Mar (1) |
Site 4 [1, 2, 3, 4, 5, 10, 12] | Apr (7), May (6), Jun (7) | |
Sphaeronychus fulvus (Baly, 1879) | Site 4 [4, 5, 9, 10, 11] | Jan (1), Feb (2), Mar (8), Jul (1), Sep (2), Oct (2) |
Syphrea sp. 1 | Site 2 [1] | Aug (2) |
Site 5 [2] | Jul (1) | |
Site 7 [1, 5, 6, 7, 8, 11] | Mar (1), Apr (1), Jun (1), Jul (6), Aug (7), Sep (1), Oct (1), Nov (1), Dec (1) | |
Syphrea sp. 2 | Site 1 [8] | Aug (1) |
Site 2 [2, 3, 5, 6, 8, 9, 10, 11, 12] | Jan (2), Feb (2), May (1), Jun (2), Jul (3), Aug (42), Sep (30), Oct (24) | |
Site 3 [1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12] | May (6), Jun (3), Jul (14), Aug (87), Sep (43), Oct (31) | |
Site 4 [1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12] | Jan (3), Feb (5), Mar (4), Apr (3), May (1), Jun (1), Jul (12), Aug (6), Sep (2), Oct (2) | |
Site 5 [5, 6, 8, 11] | Mar (1), May (3), Aug (2) | |
Site 6 [1, 2, 3, 4, 5, 8, 12] | Jan (10), Feb (19), Mar (47), Apr (34), May (6), Jun (4), Jul (6), Aug (3), Sep (3), Oct (3), Nov (2), Dec (2) | |
Systena sp. 1 | Site 7 [7] | Jul (1) |
Trichaltica scabricula (Crotch, 1873). NR | Site 7 [6, 11, 12] | May (4) |
EUMOLPINAE Hope, 1840 | ||
Tribe Eumolpini Hope, 1840 | ||
Group Iphimeites Chapuis, 1874 | ||
Brachypnoea sp. 1 | Site 2 [3,4] | Mar (2) |
Site 3 [4, 6, 8, 9, 10] | Mar (3), Apr (8), Sep (4), Oct (3) | |
Site 4 [1, 2, 7, 11, 12] | Apr (1), May (5), Jun (3), Aug (1) | |
Site 6 [3, 6, 10] | Mar (1), Apr (1), Jun (1) | |
Brachypnoea sp. 2 | Site 1 [1] | Jun (1) |
Colaspis melancholica Jacoby, 1881 | Site 2 [9] | Jun (1), Jul (1) |
Site 4 [4] | Jun (1) | |
Deuteronoda suturalis (Lefèvre, 1878)* | Site 5 | Jun |
Tribe Adoxini Baly, 1863 | ||
Group Leprotites Chapuis, 1874 | ||
Xanthonia sp. 1 | Site 2 [3] | Feb (1) |
Site 3 [5, 6, 7, 8, 9, 12] | Apr (2), May (1), Jun (3), Jul (2) | |
Site 4 [1, 2, 3, 4, 5, 6, 7, 9, 10, 11, 12] | Apr (5), May (7), Jun (6), Jul (5), Aug (2) | |
Site 5 [2, 4, 5, 6, 8, 9, 10, 11, 12] | Mar (12), Apr (13), May (22), Jun (4), Jul (1), Aug (2) | |
Site 6 [1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12] | Jan (6), Feb (12), Mar (56), Apr (59), May (37), Jun (10), Aug (8) | |
Xanthonia sp. 2 | Site 4 [4, 5, 6, 8, 9, 10, 11, 12] | Apr (12), May (7), Jun (6), Jul (1) |
Xanthonia sp. 3 | Site 3 [7, 10] | Sep (3), Oct (3) |
Site 4 [12] | Sep (1), Oct (1) | |
Xanthonia sp. 4 | Site 2 [11] | Apr (1) |
Site 3 [4, 6, 7, 10] | Mar (4), Apr (2) | |
Xanthonia sp. 5 | Site 3 [5, 12] | Jul (1), Aug (2), Sep (2), Oct (2) |
Site 4 [5, 8, 9, 11] | May (1), Jun (1), Jul (6) | |
CRYPTOCEPHALINAE Gyllenhal, 1813 | ||
Tribe Cryptocephalini Gyllenhal, 1813 | ||
Subtribe Pachybrachina Chapuis, 1874 | ||
Griburius montezuma (Suffrian, 1852) | Site 7 [4] | Jun (2) |
Griburius sp. 1 | Site 7 [1] | Jun (2) |
Pachybrachis sp. 1 | Site 1 [1, 3, 4, 5, 9, 10, 11] | Mar (1), Apr (1), Jul (1), Aug (2), Sep (4), Oct (4), Nov (1), Dec (1) |
Site 2 [3, 4, 5, 6, 8, 11] | Apr (1), Aug (3), Sep (3), Oct (3) | |
Site 4 [12] | Jul (1) | |
Site 5 [2, 4, 5] | May (2), Sep (3), Oct (3) | |
Site 6 [6, 7, 8, 9, 10, 12] | Mar (7), Apr (3), May (3), Jun (4), Jul (1), Aug (4), Sep (3), Oct (3) | |
Site 7 [1, 2, 3, 4, 5, 6, 7, 9, 11, 12] | Jan (4), Feb (5), Mar (5), Apr (1), May (1), Jun (2), Jul (3), Aug (5), Sep (5), Oct (4), Nov (3), Dec (3) | |
Pachybrachis sp. 2 | Site 2 [3, 5, 7, 9] | Mar (1), Apr (5), May (5), Aug (1) |
Pachybrachis sp. 3 | Site 5 [4] | Jul (1) |
Site 6 [12] | Jun (1) | |
Pachybrachis sp. 4 | Site 1 [3, 4, 7, 10, 11, 12] | Jan (2), Feb (2), Mar (4), May (1), Aug (1) |
Site 3 [3] | May (1) | |
Site 4 [2, 12] | Jun (1), Sep (1), Oct (1) | |
Site 7 [4, 7, 8, 9, 11, 12] | Jan (4), Feb (14), Mar (3) | |
Pachybrachis sp. 5 | Site 1 [11] | Mar (1), Aug (1) |
Site 2 [3, 4, 7, 11] | Mar (1), Jun (1), Aug (2) | |
Site 3 [3, 4, 7, 8, 10] | Mar (1), Apr (3), May (1), Jun (2), Jul (7), Aug (1) | |
Site 4 [1, 2, 7, 9, 12] | Apr (1), May (4), Jun (1), Jul (1) | |
Site 5 [4] | Apr (1) | |
Site 6 [10] | Jul (1) | |
Site 7 [12] | Aug (4) | |
Pachybrachis sp. 6 | Site 2 [7] | Sep (1), Oct (1) |
Pachybrachis sp. 7 | Site 1 [4] | Jun (1), Aug (1), Sep (2), Oct (2) |
Site 2 [3] | Sep (1), Oct (1) | |
Site 7 [2, 3, 4, 5, 7, 8, 9, 10] | Jan (2), Feb (2), Mar (1), Jun (7), Jul (5), Aug (3), Sep (6), Oct (6) | |
Pachybrachis sp. 8 | Site 1 [4, 8] | Jan (1), Feb (1), Sep (1), Oct (1) |
Site 2 [1] | Mar (1) | |
Site 3 [7] | Jul (1) | |
Site 7 [1, 3, 4, 6, 8, 9, 11, 12] | Jan (1), Feb (2), Mar (3), Jun (1), Jul (1), Aug (1), Sep (2), Oct (2), Nov (7), Dec (5) | |
Pachybrachis sp. 9 | Site 2 [3, 4, 5, 9] | Mar (2), Apr (2), May (1), Jul (1), Aug (1) |
Site 7 [3] | Apr (1) | |
Pachybrachis sp. 10 | Site 5 [1] | Jan (2), Feb (2), Aug (1) |
Pachybrachis sp. 11 | Site 5 [2] | Jan (1), Feb (1) |
Pachybrachis sp. 12 | Site 1 [4] | Sep (1), Oct (1) |
Subtribe Cryptocephalina Gyllenhal, 1813 | ||
Cryptocephalus downiei Riley & Gilbert, 2000 | Site 7 | Apr ( |
Cryptocephalus duryi Schaeffer, 1906* | Site 5 | Apr |
Cryptocephalus guttulatus Olivier, 1808 | Site 3 [11] | Jul (1) |
Site 4 [1, 8, 9, 10, 11, 12] | Jun (1), Jul (5), Aug (2) | |
Site 5 [3] | Jun (1) | |
Cryptocephalus trizonatus Suffrian, 1858 | Site 1 [1, 6, 9, 11] | Mar (1), Jun (1), Sep (1), Oct (1), Nov (2) |
Site 2 [3, 10, 12] | Aug (1), Sep (1), Oct (1), Nov (1) | |
Site 7 [1, 4, 9, 10, 11] | Apr (1), Aug (1), Nov (5), Dec (2) | |
Cryptocephalus umbonatus Schaeffer, 1906 | Site 1 [4, 11, 12] | Jul (1), Sep (2), Oct (1) |
Site 2 [3, 5, 6, 8, 9, 10, 11] | Jun (9), Jul (1), Aug (4), Sep (1), Oct (1) | |
Site 3 [3, 7, 8, 11, 12] | Jun (2), Jul (2), Sep (1), Oct (1) | |
Site 4 [4] | Aug (1) | |
Site 5 [1, 2, 12] | Aug (1), Sep (2), Oct (2) | |
Site 7 [1] | Aug (1) | |
Diachus auratus (Fabricius, 1801) | Site 1 [5, 6, 9, 10] | Mar (4) |
Site 2 [6] | Jan (1), Feb (1) | |
Site 3 [2, 10] | Mar (1), Apr (2) | |
Site 4 [2, 9, 10, 12] | Jan (1), Feb (2), Mar (5), May (1), Jun (2), Sep (1), Oct (1) | |
Site 5 [3] | Mar (2) | |
Site 6 [1] | Jan (1), Feb (1) | |
Site 7 [7] | Jan (2), Feb (3) | |
Diachus sp. 1 | Site 7 [1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 12] | Jan (10), Feb (15), Mar (19), Apr (8), May (4), Jun (6), Jul (55), Aug (40), Sep (69), Oct (50), Nov (98), Dec (44) |
Tribe Clytrini Lacordaire, 1848 | ||
Subtribe Clytrina Lacordaire, 1848 | ||
Anomoea rufifrons mutabilis (Lacordaire, 1848) | Site 3 [3] | Jul (2) |
Site 4 [1] | May (1) | |
Site 7 [3, 8] | Jun (7) | |
Smaragdina agilis (Lacordaire, 1848) | Site 4 [4] | Jun (1) |
Subtribe Megalostomina Chapuis, 1874 | ||
Coscinoptera aeneipennis (LeConte, 1858) | Site 2 [3, 7] | Jun (2) |
Coleorozena scapularis scapularis (Lacordaire, 1848) | Site 1 [4] | Apr (1) |
Site 2 [3] | Apr (1) | |
Coscinoptera tamaulipasi Medvedev, 2012 | Site [5, 9, 10, 11, 12] | Mar (4), Apr (1), May (2), Jul (1), Aug (1) |
Site 7 [2, 12] | Mar (2), May (1) | |
Megalostomis dimidiata Lacordaire, 1848 | Site 1 [10] | Mar (1) |
Subtribe Babiina Chapuis, 1874 | ||
Babia tetraspilota texana Schaeffer, 1933 | Site 1 [12] | Sep (1), Oct (1) |
Tribe Fulcidacini Jakobson, 1924 | ||
Diplacaspis prosternalis (Schaeffer, 1906) | Site 1 [2, 4, 9, 10] | Mar (2), Jul (2) |
Site 7 [2, 6, 8] | Mar (1), Jul (1), Aug (1) |