Corresponding author: Santiago Niño-Maldonado (
Academic editor: Astrid Eben
The study of biodiversity of
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. doi:
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
Study area.
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 (
Detailed position of sampling plots in Sierra de San Carlos. Cerro El Diente locality:
Detailed position of sampling plots in Sierra de San Carlos. Cerro El Diente locality:
Detailed position of sampling plots in Sierra de San Carlos. Ejido Carricitos y Tinajas locality:
Detailed position of sampling plots in Sierra de San Carlos. San Nicolás locality, Site 7. Dotted lines shows elevation curves.
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 |
|
|
550 | P1 |
|
|
772 |
P2 |
|
|
548 | P2 |
|
|
790 |
P3 |
|
|
544 | P3 |
|
|
784 |
P4 |
|
|
547 | P4 |
|
|
766 |
P5 |
|
|
535 | P5 |
|
|
773 |
P6 |
|
|
555 | P6 |
|
|
778 |
P7 |
|
|
561 | P7 |
|
|
760 |
P8 |
|
|
570 | P8 |
|
|
750 |
P9 |
|
|
571 | P9 |
|
|
743 |
P10 |
|
|
557 | P10 |
|
|
750 |
P11 |
|
|
560 | P11 |
|
|
755 |
P12 |
|
|
540 | P12 |
|
|
745 |
Site 3 – Oak forest | Site 4 – Cloud forest | ||||||
Sampling plot | Latitude | Longitude | Elevation | Sampling plot | Latitude | Longitude | Elevation |
P1 |
|
|
938 | P1 |
|
|
1077 |
P2 |
|
|
935 | P2 |
|
|
1065 |
P3 |
|
|
950 | P3 |
|
|
1070 |
P4 |
|
|
948 | P4 |
|
|
1055 |
P5 |
|
|
964 | P5 |
|
|
1085 |
P6 |
|
|
948 | P6 |
|
|
1077 |
P7 |
|
|
971 | P7 |
|
|
1093 |
P8 |
|
|
967 | P8 |
|
|
1112 |
P9 |
|
|
974 | P9 |
|
|
1109 |
P10 |
|
|
993 | P10 |
|
|
1102 |
P11 |
|
|
982 | P11 |
|
|
1086 |
P12 |
|
|
979 | P12 |
|
|
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 |
|
|
700 | P1 |
|
|
839 |
P2 |
|
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701 | P2 |
|
|
830 |
P3 |
|
|
704 | P3 |
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|
816 |
P4 |
|
|
712 | P4 |
|
|
814 |
P5 |
|
|
716 | P5 |
|
|
795 |
P6 |
|
|
720 | P6 |
|
|
813 |
P7 |
|
|
740 | P7 |
|
|
827 |
P8 |
|
|
757 | P8 |
|
|
846 |
P9 |
|
|
764 | P9 |
|
|
860 |
P10 |
|
|
763 | P10 |
|
|
866 |
P11 |
|
|
773 | P11 |
|
|
788 |
P12 |
|
|
776 | P12 |
|
|
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 |
|
|
502 |
P2 |
|
|
501 |
P3 |
|
|
500 |
P4 |
|
|
499 |
P5 |
|
|
499 |
P6 |
|
|
503 |
P7 |
|
|
508 |
P8 |
|
|
510 |
P9 |
|
|
508 |
P10 |
|
|
508 |
P11 |
|
|
502 |
P12 |
|
|
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
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 Late dry season Early rainy season Late rainy season
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
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.
Examples of leaf beetle biodiversity from Sierra de San Carlos, Mexico.
Observed and estimated species richness of
S |
Nonparametric indexes | Clench model | Completeness (%) | ||||
---|---|---|---|---|---|---|---|
Chao 1 | Jack 1 | ACE | S |
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* | |||||||
|
40 a | 44±3.74 | 48.96±2.94 | 45.88±0.5 | 47.5 | 0.026 | 81.69–90.9 |
|
49 b | 55.4±5.92 | 57.96±3.26 | 53.55±0.27 | 57.49 | 0.029 | 84.54–88.44 |
|
78 c | 91.88±8.32 | 102.9±5.69 | 94.66±1.17 | 102.17 | 0.075 | 75.8–84.89 |
|
76 c | 82.62±4.57 | 93.93±4.54 | 87.65±0.77 | 92.97 | 0.058 | 80.91–91.98 |
|
109 | 115.88±4.5 | 128.98±5.06 | 122.47±0 | 120.46 | 0.012 | 84.5–94 |
S
The most abundant subfamily in the study area was
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,
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
Mann-Whitney pairwise comparisons of chrysomelid abundance between elevational sites in Sierra de San Carlos, Mexico. Upper diagonal = Mann-Whitney U values. Lower diagonal =
S1–550 m | S2–760 m | S3–960 m | S4–1080 m | S5–730 m | S6–820 m | S7–500 m | |
---|---|---|---|---|---|---|---|
|
– | 6723 | 5344 | 6289 | 8234 | 6197 | 4675 |
|
<0.0001* | – | 8853 | 9920 | 8336 | 9735 | 8126 |
|
<0.0001* | 0.03007* | – | 9232 | 6782 | 9509 | 9634 |
|
<0.0001* | 0.52 | 0.1042 | – | 7778 | 10100 | 8530 |
|
0.001504* | 0.003257* | <0.0001* | 0.000181* | – | 7707 | 6002 |
|
<0.0001* | 0.3634 | 0.2197 | 0.748 | 0.000121* | – | 8758 |
|
<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; e
Elevational variation of abundance and diversity of
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 |
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 |
5.56 a | 21.47 b | 16.76 c | 13.98 d | 8.89 e | 13.14 d | 27.88 g |
‡S1 to S7 = Sites 1 to 7, arranged from low to high elevation. Details about the numbering of the sites are in the Materials and Methods section. † Abundance values with different letters between columns are significantly different from each other (Kruskal-Wallis, H=100.7,
According to the PERMANOVA analysis, the leaf beetle composition between sites was statistically different (SStotal=33.16; SSwithin-group=23.22; F=5.492,
Cluster analysis of chrysomelid composition by elevational site in Sierra de San Carlos, Mexico. Delimitation of groups is indicated by red dotted line.
PERMANOVA pairwise comparisons of chrysomelid composition between elevational sites in Sierra de San Carlos, Mexico. Upper diagonal = F values. Lower diagonal =
S7–500 m | S1–550 m | S5–730 m | S2–760 m | S6–820 m | S3–960 m | S4–1080 m | |
---|---|---|---|---|---|---|---|
|
– | 7.446 | 11.65 | 10.89 | 12.17 | 9.959 | 9.959 |
|
0.0001 | – | 4.59 | 3.452 | 5.391 | 4.057 | 4.692 |
|
0.0001 | 0.0001 | – | 5.13 | 2.497 | 4.229 | 4.658 |
|
0.0001 | 0.0001 | 0.0001 | – | 4.171 | 1.595 | 3.449 |
|
0.0001 | 0.0001 | 0.0182 | 0.0001 | – | 3.231 | 3.636 |
|
0.0001 | 0.0001 | 0.0001 | 0.1151 | 0.0002 | – | 2.644 |
|
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 | |
---|---|---|---|---|---|---|---|
|
– | 0.25455 | 0.14441 | 0.16639 | 0.24752 | 0.16755 | 0.16468 |
|
25.455 | – | 0.59819 | 0.31651 | 0.19005 | 0.36993 | 0.074749 |
|
14.441 | 59.819 | – | 0.37163 | 0.18434 | 0.45965 | 0.03752 |
|
16.639 | 31.651 | 37.163 | – | 0.27422 | 0.33562 | 0.050679 |
|
24.752 | 19.005 | 18.434 | 27.422 | – | 0.45432 | 0.051339 |
|
16.755 | 36.993 | 45.965 | 33.562 | 45.432 | – | 0.067524 |
|
16.468 | 7.4749 | 3.752 | 5.0679 | 5.1339 | 6.7524 | – |
Abundance and diversity values showed significant variation between seasons (H=92.29,
Historical monthly data of precipitation and temperature within Sierra de San Carlos, Mexico.
Mann-Whitney pairwise comparisons for chrysomelid abundance between seasons in Sierra de San Carlos, Mexico. Upper diagonal = Mann-Whitney U values. Lower diagonal =
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
Parameter | Dry Season | Rainy Season | ||
---|---|---|---|---|
Early ( |
Late ( |
Early ( |
Late ( |
|
|
433 a | 888 b | 690 b | 1070 c |
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 |
15.42 a | 17.01 b | 33.38 c | 21.99 d |
Abundance values with different letters between columns are significantly different from each other (Kruskal-Wallis, H=92.29,
Spearman correlation analysis for abundance and species richness of
Temperature [°C] | Precipitation [mm] | |||
---|---|---|---|---|
|
|
|
|
|
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
Cluster analysis of chrysomelid composition by month in Sierra de San Carlos, Mexico. Delimitation of groups is indicated by red dotted line.
PERMANOVA pairwise comparisons of chrysomelid composition between months in Sierra de San Carlos, Mexico. Upper diagonal = F values. Lower diagonal =
|
|
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|
|
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– | 0.964 | 0.344 | 0.063 | 0.105 | 0.312 |
|
0.076 |
|
|
|
|
|
0.191 | – | 0.799 | 0.055 | 0.093 | 0.299 |
|
0.162 |
|
|
|
|
|
1.076 | 0.629 | – | 0.666 | 0.350 | 0.115 |
|
0.179 |
|
|
|
|
|
1.668 | 1.583 | 0.812 | – | 0.939 | 0.206 |
|
0.075 |
|
|
|
|
|
1.442 | 1.407 | 1.076 | 0.493 | – | 0.587 | 0.287 | 0.163 |
|
|
|
|
|
1.147 | 1.168 | 1.416 | 1.23 | 0.902 | – | 0.467 | 0.685 | 0.083 | 0.082 |
|
|
|
1.749 | 1.607 | 1.639 | 1.563 | 1.12 | 0.995 | – | 0.879 | 0.546 | 0.479 | 0.071 | 0.078 |
|
1.566 | 1.33 | 1.34 | 1.495 | 1.264 | 0.844 | 0.638 | – | 0.739 | 0.627 |
|
|
|
1.89 | 1.713 | 1.819 | 1.973 | 1.786 | 1.459 | 0.933 | 0.709 | – | 0.962 | 0.107 | 0.088 |
|
1.89 | 1.8 | 1.988 | 2.08 | 1.831 | 1.486 | 0.986 | 0.821 | 0.041 | – | 0.078 | 0.101 |
|
1.916 | 1.809 | 1.722 | 1.718 | 1.778 | 1.929 | 1.491 | 1.864 | 1.544 | 1.624 | – | 0.972 |
|
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.
|
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|
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|
|
|
- | 0.721 | 0.487 | 0.36 | 0.398 | 0.433 | 0.298 | 0.356 | 0.295 | 0.309 | 0.230 | 0.273 |
|
72.1 | - | 0.636 | 0.347 | 0.35 | 0.367 | 0.313 | 0.390 | 0.309 | 0.318 | 0.203 | 0.215 |
|
48.7 | 63.6 | - | 0.563 | 0.482 | 0.340 | 0.319 | 0.401 | 0.353 | 0.365 | 0.161 | 0.176 |
|
36.0 | 34.7 | 56.3 | - | 0.690 | 0.403 | 0.325 | 0.294 | 0.264 | 0.291 | 0.140 | 0.176 |
|
39.8 | 35 | 48.2 | 69.0 | - | 0.509 | 0.314 | 0.267 | 0.248 | 0.262 | 0.150 | 0.144 |
|
43.3 | 36.7 | 34.0 | 40.3 | 50.9 | - | 0.367 | 0.382 | 0.331 | 0.349 | 0.161 | 0.186 |
|
29.8 | 31.3 | 31.9 | 32.5 | 31.4 | 36.7 | - | 0.584 | 0.567 | 0.580 | 0.368 | 0.353 |
|
35.6 | 39.0 | 40.1 | 29.4 | 26.7 | 38.2 | 58.4 | - | 0.573 | 0.547 | 0.259 | 0.277 |
|
29.8 | 30.9 | 35.3 | 26.4 | 24.8 | 33.1 | 56.7 | 57.3 | - | 0.894 | 0.446 | 0.368 |
|
30.9 | 31.8 | 36.5 | 29.1 | 26.2 | 34.9 | 58.0 | 54.7 | 89.4 | - | 0.422 | 0.428 |
|
23.0 | 20.3 | 16.1 | 14.0 | 15.0 | 16.1 | 36.8 | 25.9 | 44.6 | 42.2 | - | 0.698 |
|
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,
Correspondence analysis of chrysomelid abundance obtained per month at each elevational site in Sierra de San Carlos, Mexico.
The 113 species of
Although the sampled area for both studies was almost the same (33,600 m2 in Sierra de San Carlos
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
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
The species richness of
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
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
Taxon | Site [plot] | Month (abundance) |
---|---|---|
Tribe |
||
Site 4 [4] | Aug (1) | |
Site 5 [6] | Aug (1) | |
Site 3 [2] | Nov (1), Dec (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) | |
Site 5 [1] | May (1) | |
Site 3 [3, 11] | Jul (1), Sep (1), Oct (1) | |
Site 5 [3] | Aug (1) | |
Site 3 [1, 12] | Aug (1), Sep (1), Oct (1) | |
Site 4 [1, 7] | Jul (1), Aug (1) | |
Tribe |
||
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) | |
Site 7 [6] | Sep (1), Oct (1) | |
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 |
||
Site 4 [8, 9] | Jun (3), Jul (2), Aug (1) | |
Tribe |
||
Site 1 [4] | Sep (1), Oct (1) | |
Site 7 [1, 2, 12] | Jun (1), Sep (7), Oct (7), Nov (3), Dec (2) | |
Tribe |
||
Site 1 | May | |
Site 2 [5] | May (1), Jul (1) | |
Site 3 [4] | May (1) | |
Site 5 [3] | Jun (1) | |
Site 5 [7] | Aug (1) | |
Site 6 [1] | Jul (1) | |
Site 4 [8, 9, 10, 11] | May (1), Jul (3), Aug (3), Sep (4), Oct (3) | |
Tribe |
||
Subtribe |
||
Site 3 [3] | Nov (1), Dec (1) | |
Site 4 [11] | Jul (1) | |
Subtribe |
||
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) | |
Site 2 [8, 10, 11] | Jun (2), Jul (1), Aug (2) | |
Site 3 [3, 8] | Jun (2) | |
Tribe |
||
Group Coelomerites Chapuis, 1875 | ||
Site 1 [1] | Jun (1) | |
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 | ||
Site 1 [4] | Sep (1), Oct (1) | |
Site 1 [9] | Jan (1), Feb (1) | |
Tribe |
||
Site 7 [1, 6, 10] | Jun (3), Sep (3), Oct (3) | |
Site 4 [4] | Nov (1), Dec (1) | |
Tribe |
||
Subtribe |
||
Group Diabroticites Chapuis, 1875 | ||
Site 1 [5] | Sep (1), Oct (1) | |
Site 3 [5] | Sep (1), Oct (1) | |
Site 4 [4] | Aug (2) | |
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 | ||
Site 4 [6, 10, 12] | Jun (2), Aug (2) | |
Site 5 [6] | Aug (1) | |
Tribe |
||
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) | |
Site 5 [3, 7] | Aug (2) | |
Site 6 [1, 2] | Mar (3), Apr (1), May (1), Jul (1) | |
Site 4 [4] | Jun (1) | |
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) | |
Site 2 [3, 10] | Mar (1), Sep (1), Oct (1) | |
Site 3 [2] | Jan (1), Feb (1) | |
Site 4 [6, 8] | Apr (1), Sep (1), Oct (1) | |
Site 4 [10] | Jul (1) | |
Site 3 [1, 10] | Jul (1), Aug (1) | |
Site 5 [11] | Apr (1) | |
Site 7 [1, 4, 11, 12] | Aug (1), Sep (2), Oct (2), Nov (3), Dec (2) | |
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) | |
Site 2 [3, 4, 11] | Jan (1), Feb (1), Jul (1), Aug (1) | |
Site 3 [9] | May (1) | |
Site 6 [9] | May (1) | |
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) | |
Site 6 [11] | Jul (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) | |
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) | |
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) | |
Site 6 [11] | May (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) | |
Site 4 [4] | Aug (1) | |
Site 5 [1, 6] | Aug (2) | |
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) | |
Site 3 [1] | Sep (1), Oct (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) | |
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) | |
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) | |
Site 2 [9] | Mar (1) | |
Site 4 [1, 9, 12] | Jan (1), Feb (1), Mar (2) | |
Site 7 [6] | Sep (1), Oct (1) | |
Site 5 [3] | Jun (2), Aug (2) | |
Site 4 [6] | Jun (1) | |
Site 7 [6] | Aug (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) | |
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) | |
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) | |
Site 3 [4] | Sep (1), Oct (1) | |
Site 4 [1, 4] | Sep (4), Oct (2) | |
Site 5 [6] | Mar (1) | |
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) | |
Site 4 [7] | May (2) | |
Site 2 [2, 5, 7] | Jul (3), Aug (1), Sep (1), Oct (1) | |
Site 3 [1, 10, 12] | Jul (4) | |
Site 3 [3] | Aug (1) | |
Site 7 [11] | Aug (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) | |
Site 3 [3] | Mar (1) | |
Site 4 [1, 2, 3, 4, 5, 10, 12] | Apr (7), May (6), Jun (7) | |
Site 4 [4, 5, 9, 10, 11] | Jan (1), Feb (2), Mar (8), Jul (1), Sep (2), Oct (2) | |
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) | |
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) | |
Site 7 [7] | Jul (1) | |
Site 7 [6, 11, 12] | May (4) | |
Tribe |
||
Group Iphimeites Chapuis, 1874 | ||
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) | |
Site 1 [1] | Jun (1) | |
Site 2 [9] | Jun (1), Jul (1) | |
Site 4 [4] | Jun (1) | |
Site 5 | Jun | |
Tribe |
||
Group Leprotites Chapuis, 1874 | ||
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) | |
Site 4 [4, 5, 6, 8, 9, 10, 11, 12] | Apr (12), May (7), Jun (6), Jul (1) | |
Site 3 [7, 10] | Sep (3), Oct (3) | |
Site 4 [12] | Sep (1), Oct (1) | |
Site 2 [11] | Apr (1) | |
Site 3 [4, 6, 7, 10] | Mar (4), Apr (2) | |
Site 3 [5, 12] | Jul (1), Aug (2), Sep (2), Oct (2) | |
Site 4 [5, 8, 9, 11] | May (1), Jun (1), Jul (6) | |
Tribe |
||
Subtribe |
||
Site 7 [4] | Jun (2) | |
Site 7 [1] | Jun (2) | |
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) | |
Site 2 [3, 5, 7, 9] | Mar (1), Apr (5), May (5), Aug (1) | |
Site 5 [4] | Jul (1) | |
Site 6 [12] | Jun (1) | |
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) | |
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) | |
Site 2 [7] | Sep (1), Oct (1) | |
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) | |
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) | |
Site 2 [3, 4, 5, 9] | Mar (2), Apr (2), May (1), Jul (1), Aug (1) | |
Site 7 [3] | Apr (1) | |
Site 5 [1] | Jan (2), Feb (2), Aug (1) | |
Site 5 [2] | Jan (1), Feb (1) | |
Site 1 [4] | Sep (1), Oct (1) | |
Subtribe |
||
Site 7 | Apr ( |
|
Site 5 | Apr | |
Site 3 [11] | Jul (1) | |
Site 4 [1, 8, 9, 10, 11, 12] | Jun (1), Jul (5), Aug (2) | |
Site 5 [3] | Jun (1) | |
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) | |
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) | |
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) | |
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 |
||
Subtribe |
||
Site 3 [3] | Jul (2) | |
Site 4 [1] | May (1) | |
Site 7 [3, 8] | Jun (7) | |
Site 4 [4] | Jun (1) | |
Subtribe |
||
Site 2 [3, 7] | Jun (2) | |
Site 1 [4] | Apr (1) | |
Site 2 [3] | Apr (1) | |
Site [5, 9, 10, 11, 12] | Mar (4), Apr (1), May (2), Jul (1), Aug (1) | |
Site 7 [2, 12] | Mar (2), May (1) | |
Site 1 [10] | Mar (1) | |
Subtribe |
||
Site 1 [12] | Sep (1), Oct (1) | |
Tribe |
||
Site 1 [2, 4, 9, 10] | Mar (2), Jul (2) | |
Site 7 [2, 6, 8] | Mar (1), Jul (1), Aug (1) |