ZooKeys 6l is | [56 (20 | 6) A peer-reviewed open-access journal iene Macabre #ZooKeys http:/ / ZOO keys -pen soft. net Launched to accelerate biodiversity research Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational and temporal gradients in Sierra de San Carlos, Mexico Uriel Jeshua Sanchez-Reyes', Santiago Nifio-Maldonado’, Ludivina Barrientos-Lozano!, Shawn M. Clark?, Robert W. Jones* | Division de Estudios de Posgrado e Investigacién. Instituto Tecnolégico de Cd. Victoria. Boulevard Emilio Portes Gil No.1301, C.P 87010. Ciudad Victoria, Tamaulipas, México 2 Facultad de Ingenieria y Cien- cias. Universidad Auténoma de Tamaulipas. Centro Universitario Victoria. CP 87149. Victoria, Tamauli- pas, México 3 Brigham Young University, Monte L. Bean Life Science Museum, Provo, Utah 84602, U.S.A. 4 Facultad de Ciencias Naturales. Universidad Auténoma de Querétaro. Avenida de las Ciencias, s/n, 76230 Juriquilla, Querétaro, México Corresponding author: Santiago Nino-Maldonado (coliopteranino@hotmail.com) Academic editor: Astrid Eben | Received 19 June 2016 | Accepted 4 August 2016 | Published 15 August 2016 http://zoobank. org/42563154-D12F-4791-AF7C-6FA75C3E5387 Citation: Sanchez-Reyes UJ, Nifto-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: 10.3897/zookeys.611.9608 Abstract 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 faunis- tic 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 communi- ties 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 m? 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 Copyright Uriel Jeshua Sanchez-Reyes et al. 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. 12 Uriel Jeshua Sdanchez-Reyes et al. / ZooKeys 611: 11-56 (2016) 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 particu- lar 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. Keywords Biodiversity, chrysomelid beetles, ecological gradient, elevation, seasonality, sky island Introduction 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 (Jolivet et al. 2009). As a predomi- nantly phytophagous group, some species are important crop pests, while others are used efficiently to control weeds. This characteristic also makes them an important component of ecosystems, as they can compete with other herbivores (Gémez and Gonzalez-Megias 2002). Also, leaf beetles have been used as indicators of regional biodiversity and environmental quality, and for monitoring changes in natural areas (Farrell and Erwin 1988, Flowers and Hanson 2003, Kalaichelvan and Verma 2005, Linzmeier et al. 2006, Baselga and Novoa 2007, Aslan and Ayvaz 2009). Because of their importance, numerous and detailed taxonomical works on Chrysomelidae had been conducted in North America north of Mexico, Central America, and South America (Riley et al. 2003). Recently, faunistic studies have been increasing for the Mexican leaf beetle fauna. 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 (Andrews and Gilbert 2005), the central and southern portions of the country, principally at the Biosphere Reserve of Sierra de Huautla (Ordéfiez-Reséndiz and Lépez-Pérez 2009, Orddnez- Reséndiz et al. 2011, Ordéfiez-Reséndiz et al. 2015), the state of Oaxaca (Furth 2013), and the state of Morelos (Burgos-Solorio and Anaya-Rosales 2004, Nifio-Maldonado et al. 2016), where important faunistic and ecological data have been obtained. Other significant contributions have focused on the states of Jalisco (Nifio-Maldonado et al. 2014b, Sandoval-Becerra et al. 2015), Hidalgo (Martinez-Sanchez et al. 2009, 2010), and Veracruz (Deloya and Ordéfez-Reséndiz 2008), and on the Sierra Tarahumara Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 13 in Chihuahua (Furth 2009), as well as on country-wide studies of the tribe Alticini (Furth and Savini 1996, 1998, Furth 2006). To date, 2,174 species of Chrysomelidae are reported to be present in Mexico (Orddéfiez-Reséndiz et al. 2014), but the increas- ing numbers of studies have provided new distribution data, as well as species recorded for the first time in the country (Medvedev et al. 2012, Moseyko et al. 2013, Garcfa- Robledo et al. 2014, Lépez-Pérez et al. 2015, Sanchez-Reyes et al. 2015b). However, much of the faunistic information about the distribution and presence of the species in Mexico, including most recent compilations (i.e., Ordéfez-Reséndiz 2014, Nifo- Maldonado et al. 2016), are based principally on documents and studies published at least over 30 years ago (Jacoby 1880, 1881, 1882, 1883, 1884, 1885, 1886, 1887, 1888a, 1888b, 1889, 1890, 1891, 1892, Baly 1885, 1886, Champion 1893, 1894, Blackwelder 1946, Wilcox 1975, Moldenke 1970), and from collection localities cited in original descriptions of species. Moreover, many of the species from these sources lack specific localities and were only labeled as “Mexico.” This clearly demonstrates the need for new faunistic studies which provide accurate data on the present distributions of chrysomelid species in Mexico. 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 Tamauli- pas. To date, 250 species have been recorded from this state, which now ranks fourth in chrysomelid diversity from Mexico (Nifio-Maldonado et al. 2014a, Orddfiez-Reséndiz et al. 2014). Faunistic data from the state have been obtained from EI Cielo Biosphere Reserve (Nifio-Maldonado et al. 2005) and Peregrina Canyon, where detailed ecologi- cal and distributional patterns were also described (Sanchez-Reyes et al. 2014). Both areas are located within the Sierra Madre Oriental, and they are included in a protection category. Another very important area for biological conservation in Mexico and the northeastern portion of the country is the Sierra de San Carlos. This mountain range has been categorized as a terrestrial area with high to extreme priority for conservation, because of its insular geographical location within the northern gulf coastal plain and because of the relatively well-preserved nature of its natural resources (Arriaga et al. 2000, CONABIO 2007). Despite its biological interest, the only known studies from this area have focused on vegetation (Martinez 1998, Briones-Villarreal 1991), and only a few groups of insects (Meléndez-Jaramillo et al. 2014, 2015). Also, preliminary faunis- tic and ecological research on Chrysomelidae has been conducted there, showing inter- esting patterns and new species distributions (Sanchez-Reyes et al. 2015a). 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 (Ricklefs 2006, 2007). Elevation is one of the most studied gradients in species richness and diversity, because it dictates changes in environmental variables and abiotic factors such temperature, humidity, wind velocity, land area, and total ra- diation (Korner 2007, Sundqvist et al. 2013), which in turn determine distribution of species (Hodkinson 2005). The resulting patterns from these environmental influences 14 Uriel Jeshua Sdanchez-Reyes et al. / ZooKeys 611: 11-56 (2016) are different according to the studied taxa, spatial scale, or geographic region, although evidence has shown that the most common pattern is a peak in diversity at mid-eleva- tions (Lomolino 2001, Rahbek 2005, McCain and Grytnes 2010, Sanders and Rahbek 2012, Guo et al. 2013), including a peak in diversity of Chrysomelidae (Furth 2009, Sanchez-Reyes et al. 2014). Also, temporal and seasonal gradients, which determine various patterns of species diversity, are related to changes in elevation (K6rner 2007). Indeed, factors that change according to elevation, such as temperature, humidity, and vegetation, are highly variable during seasonal and temporal succession in the same mountain (Barry 2008). All these characteristics show that mountains with their elevational gradients, such as Sierra de San Carlos, can be used as key natural scenarios for analysis of patterns of diversity and also for a future assessment of changes in distri- bution of leaf beetles and other taxa, related to climate change (McCain and Grytnes 2010, Sundqvist et al. 2013). Based on these criteria, the objectives of our study were to: 1) conduct a taxonomic inventory of chrysomelid species from the Sierra de San Carlos, Tamaulipas, Mexico; 2) analyze the elevational and seasonal patterns of species richness, abundance, and diversity of this taxon in the study area; and 3) identify the effects of the interaction between sites and months on leaf beetle communities. Methods Study area The study was conducted in the Sierra de San Carlos (Figure 1), which includes the municipalities of Burgos, Cruillas, Jiménez, San Carlos, San Nicolas, and Villagran located in the central-west portion of the state of Tamaulipas, and also the municipal- ity of Linares in the extreme eastern part of the state of Nuevo Ledn, Mexico (Arriaga et al. 2000). The Sierra de San Carlos comprises an area of 2320 km’, being a polygon with the northwestern limit at 24°52.000'N, 99°12.067'W, and the southeastern limit at 24°23.050'N, 98°32.667'W. It constitutes an isolated mountain range within the Tamaulipas biogeographic province, bounded at the south by the Mexican gulf prov- ince (Morrone et al. 2002). Almost all vegetation types within the Sierra have a high conservation status, and they occur in principally temperate ecosystems (oak and pine forests) in the mountain areas, but there are also various types of tropical scrub vegeta- tion in the lower areas, as well as other vegetational communities. One of the most important characteristics of Sierra de San Carlos is its designa- tion 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 biogeographi- cal island (“sky island”) due to its isolation from other nearby mountain ranges, such as the Sierra Madre Oriental and Sierra de Tamaulipas (Arriaga et al. 2000). Accordingly, some areas with median, high and extreme conservation priority are located within Sierra de San Carlos (CONABIO 2007). Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 15 -110 -100 -90 -99°8' -98°55' _ -98°42' -110 -100 -90 99°9'0"W oF OW -99°8' = -98°55'——-9842' 98°48'0"W 98°41'0"W 98°55'0"W 24°40'0"N 24°40'0"N 24°35'0"N 24°35'0"N Zz Z S 2S S S sh p4) 30 g pa 30 = a ees Ki) omecters oA 99°9'0"W 99°2'0"W 98°55'0"W 98°48'0"W 98°41'0"W Elevation Range [ii 4s: - 545) 657 - 710} si3 - 880 1071 - 11s8{ 4} 1489 - 1799 Bi 2s: im 406 i s45 > 603 710 - 761 | ‘ 968 oe 1188 i 1323 Autor: Uriel Jeshua Sanchez-Reyes GR sr 603s A 01-1 968-071 ozs nae, Stirs Figure |. Study area. A Location of Tamaulipas in Mexico B Location of Sierra de San Carlos within Tamaulipas C Study area (red square) within Sierra de San Carlos D Details of study area: 1 = Cerro El Diente, 2 = Ejido Carricitos y Tinajas, 3 = San Nicolas. Site location We selected seven sampling sites distributed in three localities within Sierra de San Carlos (Figure 1D), including various elevations and vegetation types, as well as vari- ous conservation priorities. The first locality was Cerro El Diente which included four sampling sites: Site 1) containing submountain scrub vegetation, at a mean elevation of 550 masl; Site 2) consisting of Tamaulipan thorny scrub vegetation, at a mean eleva- tion of 760 masl; Site 3) had oak forest vegetation, at a mean elevation of 960 mas; and Site 4) with Cloud Forest vegetation, at a mean elevation of 1080 masl. The second locality was Ejido Carricitos y Tinajas which had two sampling sites. These were: Site 5) with secondary elements of riparian vegetation, at a mean elevation of 730 masl; and Site 6) containing oak and pine forest vegetation, at a mean elevation of 820 masl. The 16 Uriel Jeshua Stnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) third locality was San Nicolas with a single sampling site: Site 7) with submountain and ‘Tamaulipan thorny scrub vegetation, at a mean elevation of 500 masl. Both the Cerro El Diente and Ejido Carricitos y Tinajas are located within areas with extreme conservation priority; the locality of San Nicolas belongs to a median conservation category (CONABIO 2007). Twelve sampling plots of 400 m? each (20 x 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 (Scheiner 2003). The number of plots sampled was established by Clench analysis, with 70% as a minimum limit of completeness (for a detailed analysis, see Jiménez-Valverde and Hortal 2003). The plots were previously and randomly located within each of these sites, using GIS software. This was accom- plished by: 1) creating a polygon shape for the selected area of the sampling site, and 2) delimiting a square graticule inside this polygon using the Repeating Shapes tool, with the sides of the squares 20 meters in length; both procedures were made using ArcView GIS (1992-1999). 3) The graticule was exported to IDRISI Selva (1987-2012) and converted to a .kml format archive; 4) the new archive was displayed in Google Earth Pro software, permitting the visualization of the graticule of real dimension plots dis- played over actual satellite imagery; and finally, 5) this graticule was treated as a coor- dinate system (rows and columns), and the location of each one of the 12 plots was selected by the random numbers tool in Microsoft Office Excel. These methods were employed for the seven sites, and the only difference was the dimensions of the polygon shape of the sample area which were adjusted to the vegetational communities. Plots were georeferenced and later located in the field. When a plot was located in areas not accessible or impossible to sample (i.e. steep hills, areas of bare soil), it was moved to the closest location where vegetational cover was available. Detailed geographical data (latitude and longitude coordinates, elevation) for each plot within each site are pre- sented in Tables 1-3; detailed spatial arrangement of sampling plots within each site is presented in Figures 2—5. Collection and processing of specimens 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 Sanchez-Reyes et al. (2014). Also, leaf beetles encountered independent of the standardized sweeps were added to the species checklist. Specimens are deposited in the collection of the Facultad de Ingenieria y Ciencias at the Universi- dad Auténoma de Tamaulipas, Ciudad Victoria, Tamaulipas, Mexico. Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 17 Table |. 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 Latitude Longitude | Elevation Sanipling Latitude Longitude | Elevation plot plot Pl 24°33.020'N | 98°57.004'W 24°32.468'N | 98°57.454'W ti2 P2 24°33.048'N | 98°57.013'W 24°32.471'N | 98°57.402'W 790 P3 24°33.062'N | 98°56.960'W 24°32.471'N | 98°57.374'W 784 P4 24°33.068'N | 98°57.047'W 24°32.492'N | 98°57.353'W 766 P5 24°33.104'N | 98°57.073'W 24°32.501'N | 98°57.383'W LED P6 24°32.996'N | 98°57.096'W 24°32.490'N | 98°57.416'W 778 P7 24°32.936'N | 98°57.082'W 24°32.496'N | 98°57.460'W 760 P8 24°32.920'N | 98°57.173'W 24°32.522'N | 98°57.469'W 750 FS 24°33.031'N | 98°57.333'W 24°32.537'N | 98°57.473'W 743 P10 24°33.105'N | 98°57.316'W 24°32.531'N | 98°57.452'W 750 Pll 98°57.237W| 560 |P11 24°32.522'N | 98°57.423'W| 755 P12 98°57.137W| 540 |P12 24°32.543'N | 98°57.441'W| 745 Site 3 — Oak forest Site 4 — Cloud forest Sanpuns Latitude rg ret Latitude Longitude | Elevation plot Piet Pl | 24°32.038'N | 98°57.496'W. 98°57.557'W | 1077 P2 24°32.018'N | 98°57.466'W 24°31.795'N | 98°57.565'W | 1065 P3 24°32.021'N | 98°57.500'W 24°31.780'N | 98°57.593'W | 1070 P4 24°32.026'N | 98°57.539'W 24°31.774'N | 98°57.622'W | 1055 P5 24°31.996'N | 98°57.489'W 24°31.753'N | 98°57.631'W | 1085 P6 24°31.984'N | 98°57.456'W 24°31.760'N | 98°57.672'W | 1077 P7 24°31.973'N | 98°57.480'W 24°31.744'N | 98°57.695'W | 1093 P8 24°32.002'N | 98°57.543'W 24°31.730'N | 98°57.738'W | 1112 P9 24°31.995'N | 98°57.516'W 24°31.738'N | 98°57.783'W | 1109 P10 24°31.968'N | 98°57.527'°W 24°31.751'N | 98°57.816'W | 1102 Pll 98°57.550'W | 982 98°57.831'W | 1086 P12 98°57.577W | 979 |P12 24°31.793'N | 98°57.868'W | 1076 Taxonomic determination Identification of specimens was made using available literature on Chrysomelidae (Wilcox 1965, White 1968, Wilcox 1972, Scherer 1983, White 1993, Flowers 1996, Riley et al. 2002, Staines 2002). Additionally, material was compared with identified specimens deposited in the collection of Chrysomelidae of the Facultad de Ingenieria y Ciencias, Universidad Auténoma de Tamaulipas. However, those specimens that could not be identified to the species level were compared with other unidentified specimens and carefully grouped into morphospecies, and so the designation of “species” in this study includes both morphospecies and determined species. Taxonomical arrange- ment follows the categories employed by Riley et al. (2003), except that the subfamily Bruchinae is not included in this study. 18 Uriel Jeshua Stnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) Table 2. Sampling data in the Ejido Carricitos y Tinajas locality, Sierra de San Carlos, Mexico (coordi- nates at center of plot; elevation in meters). Ejido Carricitos y Tinajas Site 5 — Riparian and secondary vegetation Site 6 — Oak and pine forests Sampling Latitude Longitude | Elevation i Latitude Longitude | Elevation plot plot Pl 24°35.807'N | 99°2.450'W 24°35.397'N | 99°3.037'W 839 P2 24°35.789'N | 99°2.484'W 24°35.420'N | 99°3.023'W 830 PS 24°35.764'N | 99°2.508'W 24°35.440'N | 99°3.041°W 816 P4 24°35.727'N | 99°2.534'W 24°35.463'N | 99°3.017'W 814 Pp 24°35.684'N | 99°2.600'W 24°35.491'N | 99°3.028'W f95 P6 24°35.719'N | 99°2.654'W 24°35.567'N | 99°3.067'W 813 ee 24°35.673'N | 99°2.766'W 24°35.563'N | 99°3.101'W 827 P8 24°35.632'N | 99°2.851'W 24°35.575'N | 99°3.124'W 846 P9 24°35.605'N | 99°2.894'W 24°35.579'N | 99°3.146'W 860 P10 24°35.571'N | 99°2.870'W 24°35.577'N | 99°3.175'W 866 we PLL | 24°35.545'N 773 24°35.548'N 788 P12 | 24°35.533'N 776 24°35.584'N 780 Table 3. Sampling data in the San Nicolas 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 Elevation Pl 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 PA 499 P5 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 508 P10 24°32.485'N 98°46.702'W 508 Pll 24°32.334'N 98°46.922'W 502 P12 24°32.295'N 98°46.906'W 503 Organization of seasonal data We obtained environmental data from two meteorological stations located in the mu- nicipalities of San Carlos and San Nicolas 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 Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 19 98°S57'20"W 98°S7'10"W 98°57'0"W a ‘a t + fag) co laa) fae) ° ° =n a “A N vo ‘a tm ™ ne) ne NN A laa) foe) [o) °° = + ron N a a S S ne Le NN “ sf iS S st eas Sole DigitalGlobemG s = CHIBI 8, iso, USGS, INEXG ot coinapohe, batsi , IGN, IGP, Steet pe eine ie 98°57'20"W 98°57'10"W 98°57'0"W 98°57'30"W 98°57'25"W 98°57'20"W ‘a a cn on om Si “ “ ia) ioe) [e) ° =a mr ro NC a a S = sg) sg NN NN laa) ian) ° ie) a N N “N “ ioe) fon) 1e) ° + i or ro Z Z ey ; S ae ; = in }Source: Esii, DightalGlobe, Geokye, Earthstar ~----~" ie ays 20 | 0 | 0 Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Early dry season Late dry season Early rainy season Late rainy season Figure 9. Historical monthly data of precipitation and temperature within Sierra de San Carlos, Mexico. 0.0 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 Linkage Distance Figure 10. Cluster analysis of chrysomelid composition by month in Sierra de San Carlos, Mexico. Delimitation of groups is indicated by red dotted line. 34 Uriel Jeshua Stnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) NOVEMBER e e DECEMBER fal 550 730 = JANUARY ®@ SEPTEMBER [) e e FEBRUARY |.) = OCTOBER Dimension 2; Eigenvalue: 0.6060 (18.80% of Inertia) “LOD (O85 S06 3049 RO~ “0G 0.2 0.4 0.6 0.8 1.0 1.2 1.4 Dimension 1; Eigenvalue: 0.20076 (62.68% of Inertia) Figure | 1. Correspondence analysis of chrysomelid abundance obtained per month at each elevational site in Sierra de San Carlos, Mexico. Discussion Faunistic inventory and biodiversity of Chrysomelidae in Sierra de San Carlos 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 (Nifio-Maldonado et al. 2014a) and 5% of the to- tal reported Mexican leaf beetle fauna (Ordéfiez-Reséndiz et al. 2014). Also, Trichalti- ca scabricula represents a new species record for the chrysomelid fauna of Mexico, since this species was previously known only from numerous states in the United States, including Texas (Riley et al. 2003). Additionally, the high proportion of individuals identified as unnamed morphospecies in this and other studies suggests that the actual species richness of Chrysomelidae from Tamaulipas is still greater. The number of spe- cies recorded here from Sierra de San Carlos is lower when compared to other similar studies conducted with similar methods in the region. The most related study was done in Peregrina Canyon, approximately 100 km to the southwest, near Ciudad Victoria also in the state of Tamaulipas, from which 240 total samples, 2,228 specimens and 157 species were obtained (Sanchez-Reyes et al. 2014). In Sierra de San Carlos, 1,008 Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... eB) samples and 3,200 specimens were obtained, but only 109 species were recorded. Still, even with the low number of species obtained, Galerucinae was the most dominant subfamily, which is consistent with the observed patterns in various other studies (Bou- zan et al. 2015), including those conducted in other areas of Tamaulipas and in other states in Mexico (Nifio-Maldonado et al. 2005, Orddfiez-Reséndiz and Lépez-Pérez 2009, Sanchez-Reyes et al. 2014). Although the sampled area for both studies was almost the same (33,600 m? in Sierra de San Carlos vs. 37,500 m* in Peregrina Canyon), the total and site-season inventory completeness in Peregrina Canyon was close to 70% (Sanchez-Reyes et al. 2014), suggesting a higher proportion of species to be added to that inventory (Jimé- nez-Valverde and Hortal 2003), while values for this study were close to or above 90%. Moreover, seven different vegetational communities were sampled in this study, while only three sites were studied in Peregrina Canyon. These data indicate that the number of chrysomelid species present in Sierra de San Carlos was considerably less than the sites within the Sierra Madre Oriental. Because leaf beetles are dependent of their associated vegetational communities, we expected to find a higher species rich- ness at our study area. Our data suggest that the insular nature of the geographical location of Sierra de San Carlos (Arriaga et al. 2000) results in lower species richness of Chrysomelidae when compared with habitats connected to the Sierra Madre Oriental, as is the Peregrina Canyon. Also, the lesser richness of plants in Sierra de San Carlos (441 compared to 676 species, Martinez 1998, Briones-Villarreal 1991) is probably directly correlated with the smaller number of leaf beetles, when compared with Per- egrina Canyon in Altas Cumbres Natural Protected Area, where at least 1,164 species of vascular plants have been documented (Garcia-Morales et al. 2014). Contrary to species richness, the higher diversity values obtained in Sierra de San Carlos possibly reflect a lower degree of anthropogenic disturbance and a higher quality of natural resources and vegetational communities (Arriaga et al. 2000). These factors may favor a more balanced ecological process, leading to higher evenness in abundance of species and thus the higher diversity values (Magurran 2004). Elevational and seasonal effects on diversity patterns of Chrysomelidae Elevation is one of the most important factors driving ecological communities, because the abiotic factors and biotic variables together modify species richness and composi- tion of assemblages. Recent evidence suggests that the most common elevational pat- tern is the increase of diversity and species richness at intermediate elevations (Rahbek 2005, McCain and Grytnes 2010, Sanders and Rahbek 2012, Guo et al. 2013), which has been documented for various groups of Coleoptera (Escobar et al. 2005, Fernandez et al. 2010), including Chrysomelidae (Furth 2009, Sanchez-Reyes et al. 2014). Other studies have shown a decrease of species richness with increasing elevation for various groups of insects (Wolda 1987, McCoy 1990, Sanchez-Ramos et al. 1993, Araujo and Fernandes 2003, Maveety et al. 2011, Jones et al. 2012). Indeed, it has been observed that species richness, abundance and diversity of Lepidoptera decreases with increas- 36 Uriel Jeshua Sanchez-Reyes et al. / ZooKeys 611: 11-56 (2016) ing elevation in Cerro El Diente (Meléndez-Jaramillo et al. 2015), which is one of the sampling localities in our study. However, the leaf beetle communities here analyzed did not show any consistent correlation with the elevation, such as those observed in other studies of Chrysomelidae along elevational gradients (Furth 2009, Sanchez- Reyes et al. 2014, Bouzan et al. 2015). It has been determined that small elevational ranges, even with more intensive sampling (as was done in Sierra de San Carlos), could exhibit non-unimodal patterns, due to a limited geographical range over which such patterns could be expressed (Guo et al. 2013). For example, the locality of Cerro El Diente includes an elevation range from 400 to 1200 masl, but in our study only four sites were sampled at this area. It is possible that a more stepped sampling design might result in consistent patterns of species richness, abundance and diversity, as observed in other studies conducted exclusively at this locality (Meléndez-Jaramillo et al. 2015, Sanchez-Reyes et al. 2015a). However, we suggest that the insular nature of Sierra de San Carlos, its geomorphic properties, and the lack of connectivity between elevational sampling sites in different localities, could be the main drivers of results here obtained, because these factors have been recognized as key determinants of biodiversity patterns within elevational gradients (Bertuzzo et al. 2016). 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 (Jiménez-Valverde and Hortal 2003). On this basis, we affirm that 1) highest values of diversity from Sierra de San Carlos were present at the highest and lowest elevations at the Cerro El Diente locality (Site 4, Cloud forest at 1080 mas}; and Site 1, Submountain scrub at 550 masl), and also 2) the highest species richness was recorded in the highest elevation site from the same locality (Site 4, 1080 masl). This could be due to the vegetational composition and the characteristics at that area, since the contrasting more humid areas in the Cloud forest in the highest elevation site must be favoring the higher values obtained of diversity and species richness. Besides, we found higher values of species richness and diversity at sites within Cerro El Di- ente, compared with other sites from the Ejido Carricitos y Tinajas and San Nicolas localities. Land area is a determining factor in shaping communities within elevation gradients (K6rner 2007). However, this factor might not affect species richness and di- versity in our study, as fewer species were collected from more extensive sampling areas, such as Site 7 and both sites from the Ejido Carricitos y Tinajas locality; contrarily, in- termediate vegetational communities from Cerro El Diente, even with sampling plots distributed randomly within corresponding elevational intervals, covered a smaller area (Figures 2, 3) but presented higher values of species richness and diversity. Consider- ing that chrysomelid communities are directly influenced by plant-associated variables (Bach 1981, Rehounek 2002, Aslan and Ayvaz 2009, Sen and Gék 2009), we attrib- ute these results to the higher quality of the vegetational communities at the Cerro El Diente locality (Arriaga et al. 2000, CONABIO 2007). These findings highlight the significance of this locality within Sierra de San Carlos, and constitute support for its designation as an area with extreme priority for conservation. Also, the presence of Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 37 Cloud forest in Sierra de San Carlos and the associated communities of leaf beetles are remarkable, since this ecosystem has a relict distribution in Mexico and is considered as an important center for high levels of biodiversity and endemism (Gual-Diaz and Rendén-Correa 2014); thus, our results contribute to the categorization of Sierra de San Carlos as a sky island and emphasize its protection urgency. 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 (Meléndez-Jaramillo et al. 2015). It has been determined that habi- tat heterogeneity and diverse characteristics of conservation areas can promote high beta diversity or low similarity between sites, regardless of the distance (Linzmeier and Ribeiro-Costa 2009). This was observed with the lowest site of Cerro El Diente and the other three sites within the same locality, as they formed different faunistic groups, although they are geographically close to each other. Moreover, we found that the low- est elevational sites from Sierra de San Carlos (Site 1 and Site 7) formed a faunistic group even when these were from distant localities. Undoubtedly, this is due to similar vegetational communities, which lead to similar leaf beetle faunas. Conversely, both sites were very different in terms of abundance of species, thus reflecting the specific responses of each species to abiotic and biotic characteristics at each site. This evidence suggests that differences in microhabitats result in very different assemblages of spe- cies, owing to an almost entirely different plant composition and also to the climatic or abiotic variation (OWdegaard 2006). Regarding seasonal analysis, patterns observed at Sierra de San Carlos were differ- ent from those recorded in the most related study in Peregrina Canyon, Tamaulipas (Sanchez-Reyes et al. 2014), as our highest values of abundance, species richness and di- versity were recorded in the rainy season. These findings are supported by other studies, because the dominance and increased abundance of adult leaf beetles in rainy or wet sea- sons is the most common result in studies of seasonal variation of this taxon and other insects (Odegaard 2006, Furth 2009, Bouzan et al. 2015), which is highly related to the increase in plant density during this period. Besides, considering that higher seasonal peaks of abundance are associated with more marked seasons (Wolda et al. 1998), we suggest that the more seasonally dry environmental conditions at Sierra de San Carlos could be a primary factor in the species of Chrysomelidae being absent from samples or less active in the dry season, due to reduction in quality and availability of host plants, and to abiotic conditions (Medeiros and Vasconcellos-Neto 1994, Awmack and Leather 2002, Ishihara and Ohgushi 2006). Furthermore, evidence has shown that phytopha- gous insects locate temporary refugia when environmental conditions are less suitable (Janzen 1973), thus not being sampled in sweep catches and resulting in low values of abundance, species richness and diversity during dry seasons in non-refugia sites. According to cluster analysis, three groups were formed, based on faunistic similar- ity 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 38 Uriel Jeshua Stnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) months by species compositions is possibly due to differential species responses to sea- sonal variations, as their temporal niche requirements are very distinct (Linzmeier and Ribeiro-Costa 2013). Similarly, seasonal variations, monthly composition and similar- ity patterns in leaf beetles reflect different ecological and biological strategies of each species, since reproduction and adult feeding phases may be different at certain times of the year. In Chrysomelidae, this has been attributed to abiotic variation during seasons, such the photoperiod, temperature, relative humidity, and to the quality and availability of host plants (Linzmeier and Ribeiro-Costa 2008, 2013). Also, seasonal and monthly patterns could reflect an existing sequence between Chrysomelidae and other phytophagous species to avoid competition and allow optimum resource exploi- tation (Pérez-Barroeta and Gurrea-Sanz 1994). 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 vari- ables (Bach 1981, Erelli et al. 1998, Rehounek 2002, Aslan and Ayvaz 2009, Sen and Gok 2009). So, although not investigated here, the study of relationships between Chrysomelidae and their associated vegetational communities is very important to understand the distribution patterns of this taxon. However, it has been noted that the interplay between the biotic and abiotic environment shapes consumer diversity along elevational gradients (Sundqvist et al. 2013). Therefore, changes in abundance, species richness and diversity within each locality, site and season/month could be driven by other factors related to elevation change, such as climate variables or abiotic environ- ment (temperature, humidity, K6rner 2007), which could be influencing the activity patterns of Chrysomelidae and the capacity of each species to obtain resources (Flinte et al. 2011, Bouzan et al. 2015). This influence of climate has been demonstrated in this study, since precipitation and temperature were significantly correlated with spe- cies richness; other studies on Chrysomelidae have reported similar effects (Linzmeier and Ribeiro-Costa 2008, Sanchez-Reyes et al. 2014). 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 environ- mental (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 (Hodkinson 2005, Sundqvist et al. 2013, Guo et al. 2013), future research on Chrysomelidae in Mexico needs to be targeted at the specific species patterns and their relationships with environmental variation, as well as at specific interactions between leaf beetle species and other eco- system components. Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... a9 Conclusions The species richness of Chrysomelidae in Sierra de San Carlos was not as high as ex- pected 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 veg- etational 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 geomor- phic characteristics of Sierra de San Carlos on the Chrysomelidae distribution. Rainy season was associated with higher values of the ecological parameters of Chrysomeli- dae, 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 assem- blages. 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 pre- cipitation with species richness. These differences and variations in faunistic composi- tion 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 require- ments 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 bio- geographical relationships of chrysomelid species existent within this and others areas in the country. Acknowledgments We are grateful to our work crew of the 2012-2014 period, which efficiently assisted in the sampling work: Edmar Meléndez-Jaramillo, Nabil Yessenia Martinez-Ruiz and 40 Uriel Jeshua Stnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) Brenda Villanueva-Alanis. Also, we thank Vannia del Carmen Gémez-Moreno, Geo- vany de Jestis Fernandez-Azuara, and Luis 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 plan- ning phases of this study, at Instituto Tecnoldgico de Ciudad Victoria. The active authorities in 2013 from San Carlos and San Nicolas municipalities granted us permission for fieldwork in different areas within Sierra de San Carlos. We are indebted to Jestis 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 Gonzalez, 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 Tecnologia (CONACYT), for a scholarship award granted for M.S. studies at the Instituto Tecnoldgico de Ciudad Victoria. The authors also thank PROMEP, for additional financial support. References Andrews FG, Gilbert AJ (2005) A preliminary annotated checklist and evaluation of the diver- sity of the Chrysomelidae (Coleoptera) of the Baja California peninsula, Mexico. Insecta Mundi 19(1—2): 89-116. Araujo LM, Fernandes GW (2003) Altitudinal patterns in a tropical ant assemblage and varia- tion in species richness between habitats. Lundiana 4(2): 103-109. ArcView GIS Version 3.2 (1992-1999) ArcView GIS Version 3.2. Environmental Systems Research Institute (ESRI), Redlands, CA. Arriaga L, Espinoza JM, Aguilar C, Martinez E, Gomez L, Loa E (2000) Regiones terrestres prioritarias de México. Comisién Nacional para el Conocimiento y uso de la Biodiversidad, México. http://www.conabio.gob.mx/conocimiento/regionalizacion/doctos/terrestres.html Aslan EG, Ayvaz Y (2009) Diversity of Alticinae (Coleoptera, Chrysomelidae) in Kasnak Oak Forest Nature Reserve, Isparta, Turkey. Turkish Journal of Zoology 33(3): 251-262. doi: 10.3906/zo00-0806-2 Awmack CS, Leather SR (2002) Host plant quality and fecundity in herbivorous insects. An- nual Review of Entomology 47: 817-844. doi: 10.1146/annurev.ento.47.091201.145300 Bach CE (1981) Host plant growth form and diversity: effects on abundance and feeding pref- erence of a specialist herbivore, Acalymma vittata (Coleoptera: Chrysomelidae). Oecologia 50(3): 370-375. doi: 10.1007/bf00344978 Baly JS (1885) Biologia Centrali-Americana, Insecta, Coleoptera, Hispidae. Vol. VI. Part 2. R. H. Porter, London, 1—72. Baly JS (1886) Biologia Centrali-Americana, Insecta, Coleoptera, Hispidae and supplement. Vol. VI. Part 2. R. H. Porter, London, 73-124. doi: 10.1017/CBO978051 1754753 Barry RG (2008) Mountain Weather and Climate Third Edition. Cambridge University Press, New York, 506 pp. Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 4] Baselga A, Novoa F (2007) Diversity of Chrysomelidae (Coleoptera) at a mountain range in the limit of the Eurosiberian region, northwest Spain: species richness and beta diversity. Entomologica Fennica 18(2): 65-73. Beh EJ (2004) Simple correspondence analysis: a bibliographic review. International Statistical Review 72(2): 257-284. doi: 10.1111/j.1751-5823.2004.tb00236.x Bertuzzo E, Carrara F, Mari L, Altermatt FE, Rodriguez-Iturbe I, Rinaldo A (2016) Geomorphic controls on elevational gradients of species richness. Proceedings of the National Academy of Sciences 113(7): 1737-1742. doi: 10.1073/pnas.1518922113 Blackwelder RE (1946) Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 4. Bulletin of the United States National Museum 185: 551-763. doi: 10.1093/aesa/39.3.543 Bouzan A, Flinte V, Macedo MV, Monteiro RF (2015) Elevation and temporal distributions of Chrysomelidae in southeast Brazil with emphasis on the Galerucinae. ZooKeys 547: 103-117. doi: 10.3897/zookeys.547.9723 Briones-Villarreal OL (1991) Sobre la flora, vegetacidn y fitogeografia de la Sierra de San Car- los, Tamaulipas. Acta Botanica Mexicana 16: 15-43. Burgos-Solorio A, Anaya-Rosales S (2004) Los crisomelinos (Coleoptera: Chrysomelidae: Chrysomelinae) del estado de Morelos. Acta Zoolégica Mexicana (NS) 20(3): 39-66. Champion CG (1893) Biologia Centrali-Americana, Insecta, Coleoptera, Cassididae. Vol. VI. Part 2. R. H. Porter, London, 125-164. Champion CG (1894) Biologia Centrali-Americana, Insecta, Coleoptera, Cassididae and sup- plement, appendix to Hispidae. Vol. VI. Part 2. R. H. Porter, London, 165-242. Colwell RK (2013) EstimateS: Statistical estimation of species richness and shared species from samples. Version 9. User’s Guide and application. http://purl.oclc.org/estimates CONABIO (2007) CONABIO, CONANP, TNC, Pronatura. Sitios prioritarios terrestres para la conservacién de la biodiversidad. Escala 1: 1000000. Comisién Nacional para el Cono- cimiento y Uso de la Biodiversidad, Comisién Nacional de Areas Naturales Protegidas, The Nature Conservancy - Programa México, Pronatura, México, D.F. Deloya CA, Ordéfez-Reséndiz MM (2008) Escarabajos (Insecta: Coleoptera). In: Manson RH, Hernandez-Ortiz V, Gallina S, Mehltreter K (Eds) Agroecosistemas cafetaleros de Veracruz: biodiversidad, manejo y conservacién. Instituto Nacional de Ecologia A. C., México, 123-134. Erelli MC, Ayres MP, Eaton GK (1998) Altitudinal patterns in host suitability for forest insects. Oecologia 117(1): 133-142. doi: 10.1007/s004420050641 Escobar F, Lobo JM, Halffter G (2005) Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes. Global Ecology and Biogeography 14(4): 327-337. doi: 10.1111/j.1466-822X.2005.00161.x Farrell BD, Erwin TL (1988) Leaf-beetle community structure in an amazonian rainforest canopy. In: Jolivet P, Petitpierre E, Hsiao TH (Eds) Biology of Chrysomelidae. Kluwer Academic Publishers, Dordrecht, 73—90. doi: 10.1007/978-94-009-3105-3_5 Fernandez V, Gamarra P, Outerelo R, Cifridn B, Baz A (2010) Distribucién de estafilininos necrofilos (Coleoptera, Staphylinidae, Staphylininae) a lo largo de un gradiente altitudinal 42 Uriel Jeshua Sanchez-Reyes et al. / ZooKeys 611: 11-56 (2016) en la Sierra de Guadarrama, Espafia. Boletin de la Real Sociedad Espafiola de Historia Natural (Seccién Bioldgica) 104(1—4): 61-86. Flinte V, Freitas S, Macedo MV, Monteiro RF (2011) Altitudinal and temporal distribution of Plagiometriona Spaeth, 1899 (Coleoptera, Chrysomelidae, Cassidinae) in a tropical forest in southeast Brazil. ZooKeys 157: 15-31. doi: 10.3897/zookeys.157.1179 Flowers RW (1996) La subfamilia Eumolpinae (Coleoptera: Chrysomelidae) en América Cen- tral. Publicacién especial de la Revista de Biologia Tropical 2: 1-60. Flowers RW, Hanson PE (2003) Leaf beetle (Coleoptera: Chrysomelidae) diversity in eight Costa Rican habitats. In: Furth DG (Ed.) Special topics in leaf beetle biology Proceedings of the 5th International Symposium on the Chrysomelidae, Iguassu Falls (Brazil), August 2000. Pensoft Publishers, Sofia-Moscow, 25—51. Furth DG (2006) The current status of knowledge of the Alticinae of Mexico (Coleoptera: Chrysomelidae). Bonner zoologische Beitrage 54(4): 209-237. Furth DG (2009) Flea beetle diversity of the Sierra Tarahumara, Copper Canyon, Mexico (Chrysomelidae: Alticinae). In: Jolivet P, Santiago-Blay J, Schmitt M (Eds) Chrysomelidae, Volume 2. Koninklijke Brill, Leiden, 131-151. doi: 10.1163/ej.9789004169470.1-299.45 Furth DG (2013) Diversity of Alticinae in Oaxaca, Mexico: A preliminary study (Coleoptera, Chrysomelidae). ZooKeys 332: 1-32. doi: 10.3897/zookeys.332.4790 Furth DG, Savini V (1996) Checklist of the Alticinae of Central America, including Mexico (Coleoptera: Chrysomelidae). Insecta Mundi 10(1—4): 45-68. Furth DG, Savini V (1998) Corrections, clarifications, and additions to the 1996 checklist of the Alticinae of Central America, including Mexico (Coleoptera: Chrysomelidae). Insecta Mundi 12(1—2): 133-138. Garcia-Morales LJ, Estrada-Castill6n AE, Garcia-Jiménez J, Villarreal-Quintanilla JA, Cantu- Ayala C, Jurado-Ybarra E, Vargas-Vazquez VA (2014) Floristica y vegetacién del Area Nat- ural Protegida Altas Cumbres, Tamaulipas, México. In: Correa-Sandoval A, Horta-Vega JV, Garcia-Jiménez J, Barrientos-Lozano L (Eds) Biodiversidad tamaulipeca, Volumen 2, Numero 1. Tecnolégico Nacional de México, Instituto Tecnolégico de Ciudad Victoria, Tamaulipas, 15-73. Garcia-Robledo C, Kuprewicz EK, Bordignon L, Bahia TDO, Rodriguez-Morales D, Aguilar- Argiiello S, Hernandez-Lara C, Puga-Ayala L, Angel-Pifa CD, Abella-Medrano A, Mar- tinez-Adriano CA, Delgado-Carrillo O, Garcia-Conejo M, Leén-Solano H, Navarrete- Jiménez A, Bastidas-Martinez A, Martinez-Roldan H, Cuyos-Palacios M, Salinas-Hijar I, Aguirre A, Hernandez-Ortiz V (2014) First record of the genus Chelobasis Gray, 1832 (Coleoptera: Chrysomelidae: Cassidinae) in Mexico (North America). The Coleopterists Bulletin 68(4): 774-775. doi: 10.1649/0010-065x-68.4.774 Gomez JM, Gonzdlez-Megias A (2002) Asymmetrical interactions between ungulates and phytophagous insects: being different matters. Ecology 83(1): 203-211. doi: 10.1890/0012-9658(2002)083[0203:AIBUAP]2.0.CO;2 Gotelli NJ, Colwell RK (2011) Estimating species richness. In: Magurran AE, McGill BJ (Eds) Biological diversity: frontiers in measurement and assessment. Oxford University Press, Oxford, 39-54. Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 43 Gual-Diaz M, Rendén-Correa A (2014) Bosques mesdéfilos de montafia de México, diversidad, ecologia y manejo. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO), México, D. E, 352 pp. Guo Q, Kelt DA, Sun Z, Liu H, Hu L, Ren H, Wen J (2013) Global variation in elevational diversity patterns. Scientific Reports 3(3007): 1-7. doi: 10.1038/srep03007 Hammer @, Harper DAT, Ryan PD (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4(1): 1-9. Hodkinson ID (2005) Terrestrial insects along elevation gradients: species and community re- sponses to altitude. Biological Reviews 80(3): 489-513. doi: 10.1017/s1464793 105006767 Hortal J, Borges PAV, Gaspar C (2006) Evaluating the performance of species richness esti- mators: sensitivity to sample grain size. Journal of Animal Ecology 75(1): 274-287. doi: 10.1111/j.1365-2656.2006.01048.x IDRISI Selva (1987-2012) IDRISI Selva 17.0. Clark Labs, Clark University, Worcester, MA. Ishihara M, Ohgushi T (2006) Reproductive inactivity and prolonged developmental time induced by seasonal decline in host plant quality in the willow leaf beetle Plagiodera ver- sicolora (Coleoptera: Chrysomelidae). Environmental Entomology 35(2): 524-530. doi: 10.1603/0046-225x-35.2.524 Jacoby M (1880) Biologia Centrali-Americana, Insecta, Coleoptera, Sagridae, Crioeridae, Meg- alopodidae, Clythridae, Cryptocephalidae. Vol. VI. Part 1. R. H. Porter, London, 1-72. Jacoby M (1881) Biologia Centrali-Americana, Insecta, Coleoptera, Cryptocephalidae, Chlamydidae, Lamprosomatidae, Eumolpidae. Vol. VI. Part 1. R. H. Porter, London, 73-144. Jacoby M (1882) Biologia Centrali-Americana, Insecta, Coleoptera, Eumolpidae, Chrysomeli- dae. Vol. VI. Part 1. R. H. Porter, London, 145-224. Jacoby M (1883) Biologia Centrali-Americana, Insecta, Coleoptera, Chrysomelidae. Vol. VI. Part 1. R. H. Porter, London, 225-264. Jacoby M (1884) Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae. Vol. VI. Part 1. R. H. Porter, London, 265-336. Jacoby M (1885) Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae. Vol. VI. Part 1. R. H. Porter, London, 337-408. Jacoby M (1886) Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae. Vol. VI. Part 1. R. H. Porter, London, 409-496. Jacoby M (1887) Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae. Vol. VI. Part 1. R. H. Porter, London, 497-584. Jacoby M (1888a) Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae. Vol. VI. Part 1. R. H. Porter, London, 585-625. Jacoby M (1888b) Biologia Centrali-Americana, Insecta, Coleoptera, Supplement to Phy- tophaga. Vol. VI. Part 1. R. H. Porter, London, 1-80. Jacoby M (1889) Biologia Centrali-Americana, Insecta, Coleoptera, Supplement to Phytopha- ga. Vol. VI. Part 1. R. H. Porter, London, 81-168. Jacoby M (1890) Biologia Centrali-Americana, Insecta, Coleoptera, Supplement to Phytopha- ga. Vol. VI. Part 1. R. H. Porter, London, 169-232. 44 Uriel Jeshua Sdnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) Jacoby M (1891) Biologia Centrali-Americana, Insecta, Coleoptera, Supplement to Phytopha- ga. Vol. VI. Part 1. R. H. Porter, London, 233-312. Jacoby M (1892) Biologia Centrali-Americana, Insecta, Coleoptera, Supplement to Phytopha- ga. Vol. VI. Part 1. R. H. Porter, London, 313-348. Janzen DH (1973) Sweep samples of tropical foliage insects: effects of seasons, vegetation types, elevation, time of day, and insularity. Ecology 54(3): 687-708. doi: 10.2307/1935359 Jiménez-Valverde A, Hortal J (2003) Las curvas de acumulacion de especies y la necesidad de evaluar la calidad de los inventarios bioldgicos. Revista Ibérica de Aracnologia 8: 151-161. Jolivet P, Santiago-Blay J, Schmitt M (2009) Research on Chrysomelidae, Volume 2. Brill, Leiden, The Netherlands, 300 pp. Jones RW, Nifio-Maldonado S, Luna-Cozar J (2012) Diversity and biogeographic affinities of Apionidae (Coleoptera: Curculionoidea) along an altitudinal gradient in El Cielo Bio- sphere Reserve of northeastern Mexico. Revista Mexicana de Biodiversidad 83: 100-109. Jost L (2006) Entropy and diversity. Oikos 113(2): 363-375. doi: 10.1111/j.2006.0030- 1299.14714.x Jost L (2007) Partitioning diversity into independent alpha and beta components. Ecology 88(10): 2427-2439. doi: 10.1890/06-1736.1 Kalaichelvan T, Verma KK (2005) Checklist of leaf beetles (Coleoptera: Chrysomeli- dae) of Bhilai-Durg. Zoos’ Print Journal 20(4): 1838-1842. doi: 10.11609/jott. zpj.1114.1838-42 Korner C (2007) The use of ‘altitude’ in ecological research. Trends in Ecology & Evolution 22(11): 569-574. doi: 10.1016/j.tree.2007.09.006 Linzmeier AM, Ribeiro-Costa CS (2008) Seasonality and temporal structuration of Alti- cini community (Coleoptera, Chrysomelidae, Galerucinae) in the Araucaria Forest of Parana, Brazil. Revista Brasileira de Entomologia 52(2): 289-295. doi: 10.1590/s0085- 56262008000200009 Linzmeier AM, Ribeiro-Costa CS (2009) Spatio-temporal dynamics of Alticini (Coleoptera, Chrysomelidae) in a fragment of Araucaria Forest in the state of Parana, Brazil. Revista Brasileira de Entomologia 53(2): 294-299. doi: 10.1590/s0085-56262009000200013 Linzmeier AM, Ribeiro-Costa CS (2013) Seasonal pattern of Chrysomelidae (Coleoptera) in the state of Parana, southern Brazil. Biota Neotropica 13(1): 153-162. doi: 10.1590/ s1676-06032013000100018 Linzmeier AM, Ribeiro-Costa CS, Marinoni RC (2006) Fauna de Alticini (Newman) (Co- leoptera, Chrysomelidae, Galerucinae) em diferentes estagios sucessionais na Floresta com Araucaria do Parana, Brasil: diversidade e estimativa de riqueza de espécies. Revista Bra- sileira de Entomologia 50(1): 101-109. doi: 10.1590/S0085-56262006000100015 Lomolino MV (2001) Elevation gradients of species-density: historical and prospec- tive views. Global Ecology and Biogeography 10(1): 3-13. doi: 10.1046/j.1466- $22%,2.001 00229 Lépez-Pérez S, Martinez-Falcén AP, Benitez-Malvido J (2015) First record of the Tribe Hemi- sphaerotini Monros and Viena (Chrysomelidae: Cassidinae) in Mexico. Southwestern En- tomologist 40(1): 241-244. doi: 10.3958/059.040.0125 Magurran AE (2004) Measuring biological diversity. Blackwell Science Ltd., Oxford, 256 pp. Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 45 Martinez M (1998) Inventario floristico de la Sierra de San Carlos, Tamps. Informe final SNIB-CONABIO proyecto No. P024. Universidad Auténoma de Tamaulipas. Instituto de Ecologia Aplicada, México, D.F, 55 pp. Martinez-Sanchez I, Nifio-Maldonado S, Barrientos-Lozano L, Horta-Vega JV (2009) Dinami- ca poblacional de Chrysomelidae (Coleoptera) en un gradiente altitudinal en tres muni- cipios del estado de Hidalgo, México. TecnoINTELECTO 6(2): 2-4. Martinez-Sanchez I, Nifio-Maldonado S, Carreén-Pérez A, Horta-Vega JV (2010) Nuevos reg- istros de Cassidinae (Coleoptera: Chrysomelidae) para el Estado de Hidalgo, México. In: Cruz-Miranda SG, Tello-Flores J, Mendoza-Estrada A, Morales-Moreno A (Eds) Ento- mologia mexicana Vol 9. Sociedad Mexicana de Entomologia A. C., Texcoco, Estado de México, 921-924. Maveety SA, Browne RA, Erwin TL (2011) Carabidae diversity along an altitudinal gradi- ent in a Peruvian cloud forest (Coleoptera). ZooKeys 147: 651-666. doi: 10.3897/zook- eys. 147.2047 McCain CM, Grytnes J-A (2010) Elevational gradients in species richness. In: Encyclope- dia of Life Sciences (eLS). John Wiley & Sons Ltd., Chichester. http://www.els.net, doi: 10.1002/9780470015902.a0022548 McCoy ED (1990) The distribution of insects along elevational gradients. Oikos 58(3): 313- 332. doi: 10.2307/3545222 Medeiros L, Vasconcellos-Neto J (1994) Host plants and seasonal abundance patterns of some Brazilian Chrysomelidae. In: Jolivet PH, Cox ML, Petitpierre E (Eds) Novel aspects of the biology of Chrysomelidae. Springer Science+Business Media, Dordrecht, Netherlands, 185-189. doi: 10.1007/978-94-011-1781-4_11 Medvedev LN, Nifio-Maldonado S, Sanchez-Reyes UJ, Moseyko AG (2012) To the knowledge of Mexican Clytrini (Chrysomelidae, Cryptocephalinae), with description of two new spe- cies of the genus Coscinoptera Lacordaire, 1848. Zoosystematica Rossica 21(2): 244-253. Meléndez-Jaramillo E, Garcia-Jiménez J, Cantt-Ayala C (2015) Pieridae (Lepidoptera: Rho- palocera) del Cerro Bufa El Diente, municipio de San Carlos, Tamaulipas, México. Ento- mologia Mexicana 2: 863-869. Meléndez-Jaramillo E, Garcia-Jiménez J, Garcia-Morales LJ, Vargas-Vazquez VA (2014) Escara- bajos copréfagos (Coleoptera: Scarabaeidae) del matorral submontano del Cerro Bufa El Di- ente, municipio de San Carlos, Tamaulipas, México. Entomologia Mexicana 1: 1201-1206. Moldenke AR (1970) A revision of the Clytrinae of North America north of the Isthmus of Panama. Stanford University, Stanford, California, 310 pp. Morrone JJ, Espinosa-Organista D, Llorente-Bousquets J (2002) Mexican biogeographic prov- inces: preliminary scheme, general characterizations, and synonymies. Acta Zooldgica Mexicana (NS) 85: 83-108. Moseyko AG, Nifio-Maldonado S, Ruiz-Cancino E, Coronado-Blanco JM (2013) A new spe- cies of the genus Rhabdopterus Lefevre, 1885 (Coleoptera: Chrysomelidae) from the state of Tamaulipas, Mexico. Russian Entomological Journal 22(2): 127-130. Nifio-Maldonado S, Riley EG, Furth DG, Jones RW (2005) Coleoptera: Chrysomelidae. In: Sanchez-Ramos G, Reyes-Castillo P, Dirzo R (Eds) Historia Natural de la Reserva de la Bids- fera El Cielo, Tamaulipas, México. Universidad Auténoma de Tamaulipas, Victoria, 417-425. 46 Uriel Jeshua Sanchez-Reyes et al. / ZooKeys 611: 11-56 (2016) Nifio-Maldonado S, Romero-Napoles J, Sanchez-Reyes UJ, Jones RW, De-Leén-Gonzalez EI (2014a) Inventario preliminar de Chrysomelidae (Coleoptera) de Tamaulipas, México. In: Correa-Sandoval A, Horta-Vega JV, Garcia-Jiménez J, Barrientos-Lozano L (Eds) Biodi- versidad Tamaulipeca, Volumen 2, Numero 2. Tecnoldgico Nacional de México, Instituto Tecnoldgico de Cd. Victoria, Tamaulipas, México, 121-132. Nifio-Maldonado S, Sanchez-Reyes UJ, Clark SM, Toledo-Hernandez VH, Corona-Lépez AM, Jones RW (2016) Checklist of leaf beetles (Coleoptera: Chrysomelidae) from the state of Morelos, Mexico. Zootaxa 4088(1): 91-111. doi: 10.11646/zootaxa.4088.1.4 Nifio-Maldonado S, Sanchez-Reyes UJ, Meléndez-Jaramillo E, Gdmez-Moreno VdC, Navar- rete-Heredia JL (2014b) Coledépteros Chrysomelidae. In: Guerrero S, Navarrete-Heredia JL, Contreras-Rodriguez SH (Eds) Biodiversidad del Estero El Salado. Universidad de Guadalajara, Guadalajara, 85-98. Odegaard F (2006) Host specificity, alpha- and beta-diversity of phytophagous beetles in two tropical forests in Panama. Biodiversity and Conservation 15(1): 83-105. doi: 10.1007/ s10531-004-3106-5 Orddfez-Reséndiz MM (2014) Catalogo de autoridades taxondomicas y base de datos curato- rial de la Familia Chrysomelidae en México. Informe final, SNIB-CONABIO. Proyecto No. HS003. Universidad Nacional Auténoma de México. Facultad de Estudios Superiores Zaragoza, México, D. F, 103 pp. Ordofiez-Reséndiz MM, Lopez-Pérez S (2009) Crisomélidos (Coleoptera: Chrysomelidae) de las Sierras de Taxco-Huautla, México. In: Estrada-Venegas EG, Equihua-Martinez A, Chaires-Grijalva MP, Acufia-Soto JA, Padilla-Ramirez JR, Mendoza-Estrada A (Eds) En- tomologia mexicana Vol 8. Sociedad Mexicana de Entomologia A. C., Texcoco, Estado de México, 946-951. Ordéfiez-Reséndiz MM, Lépez-Pérez S, Rodriguez-Mir6én G (2011) Chrysomelidae (Coleop- tera) en la Sierra Nevada, México. In: Cruz-Miranda SG, Tello-Flores J, Mendoza-Estrada A, Morales-Moreno A (Eds) Entomologia mexicana Vol 10. Sociedad Mexicana de Ento- mologia A. C., Texcoco, Estado de México, 779-784. Ordéfez-Reséndiz MM, Lopez-Pérez S, Rodriguez-Mir6n G (2014) Biodiversidad de Chrysomelidae (Coleoptera) en México. Revista Mexicana de Biodiversidad Supl. 85: 271-278. doi: 10.7550/rmb.31424 Ordofiez-Reséndiz MM, Serrano-Resendiz V, Hernandez-Sosa L (2015) Riqueza de la familia Chrysomelidae (Coleoptera) en comunidades vegetales de las Sierras de Taxco-Huautla. Entomologia Mexicana 2: 601-607. Pérez-Barroeta F, Gurrea-Sanz P (1994) Diversidad de Curculionoidea y Chrysomelidae (Co- leoptera) en prados del Pirineo central (Huesca). Lucas Mallada 6: 131-141. Rahbek C (2005) ‘The role of spatial scale and the perception of large-scale species-richness pat- terns. Ecology Letters 8(2): 224-239. doi: 10.1111/j.1461-0248.2004.00701.x Rehounek J (2002) Comparative study of the leaf beetles (Coleoptera: Chrysomelidae) in cho- sen localities in the district of Nymburk. Acta Universitatis Palackianae Olomucensis Fac- ultas Rerum Naturalium Biologica 39-40: 123-130. Ricklefs RE (2006) Evolutionary diversification and the origin of the diversity—environment relationship. Ecology 87: 3-13. doi: 10.1890/0012-9658(2006)87[3:edatoo]2.0.co;2 Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 47 Ricklefs RE (2007) History and diversity: explorations at the intersection of ecology and evolu- tion. The American Naturalist 170: 56—70. doi: 10.1086/519402 Riley EG, Clark SM, Flowers RW, Gilbert AJ (2002) Chrysomelidae Latreille 1802. In: Arnett RH, Thomas MC, Skelley PE, Frank JH (Eds) Volume 2, American Beetles, Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press LLC, Boca Raton, 617-691. Riley EG, Clark SM, Seeno TN (2003) Catalog of the leaf beetles of America north of Mexico. Coleopterists Society, special publication no. 1, 290 pp. Sanchez-Ramos G, Lobo J, Lara-Villal6n M, Reyes-Castillo P (1993) Distribucion altitudinal y estacional de la entomofauna necréfila en la Reserva de la Bidésfera “El Cielo”, Tamaulipas, México. BIOTAM 5(1): 13-24. Sanchez-Reyes UJ, Nifio-Maldonado S, Jones RW (2014) Diversity and altitudinal distribution of Chrysomelidae (Coleoptera) in Peregrina Canyon, Tamaulipas, Mexico. ZooKeys 417: 103-132. doi: 10.3897/zookeys.417.7551 Sanchez-Reyes UJ, Nifio-Maldonado S, Meléndez-Jaramillo E, Gémez-Moreno VdC, Banda- Hernandez JE (2015a) Riqueza de Chrysomelidae (Coleoptera) en el Cerro El Diente, San Carlos, Tamaulipas, México. Acta Zoolégica Mexicana (NS) 31(1): 10-22. Sanchez-Reyes UJ, Nifto-Maldonado S, Meléndez-Jaramillo E, Gémez-Moreno VdC, Vil- lanueva-Alanis B, Martinez-Ruiz NY (2015b) Primer registro de Cryptocephalus downiei Riley & Gilbert, 1999 (Coleoptera: Chrysomelidae, Cryptocephalinae) en México. Dug- esiana 22(1): 55-57. Sanders NJ, Rahbek C (2012) The patterns and causes of elevational diversity gradients. Ecog- raphy 35(1): 1-3. doi: 10.1111/j.1600-0587.2011.07338.x Sandoval-Becerra FM, Sanchez-Reyes UJ, Nifto-Maldonado S, Vasquez-Bolafios M, Herrera- Herrera JR (2015) Distribucién potencial de Helocassis Spaeth, 1952 (Chrysomelidae: Cas- sidinae) en México. Dugesiana 22(2): 121-131. Scheiner SM (2003) Six types of species-area curves. Global Ecology and Biogeography 12(6): 441-447. doi: 10.1046/j.1466-822x.2003.00061.x Scherer G (1983) Diagnostic key for the Neotropical Alticinae genera. Entomologische Arbe- iten aus dem Museum G Frey 31-32: 2-89. Sen I, Gok A (2009) Leaf beetle communities (Coleoptera: Chrysomelidae) of two mixed forest ecosystems dominated by pine—oak—hawthorn in Isparta province, Turkey. Annales Zoo- logici Fennici 46(3): 217-232. doi: 10.5735/086.046.0306 Staines CL (2002) The New World tribes and genera of Hispines (Coleoptera: Chrysomeli- dae: Cassidinae). Proceedings of the Entomological Society of Washington 104(3): 721-784. StatSoft Inc. (2007) STATISTICA, Version 8.0. http://www.statsoft.com Sundgqvist MK, Sanders NJ, Wardle DA (2013) Community and ecosystem responses to el- evational gradients: Processes, mechanisms, and insights for global change. Annual Re- view of Ecology, Evolution, and Systematics 44: 261-280. doi: 10.1146/annurev-ecol- sys-110512-135750 White RE (1968) A review of the genus Cryptocephalus in America north of Mexico (Chrysomel- idae: Coleoptera). United States National Museum Bulletin 290: 1-124. doi: 10.5479/ si.03629236.290.1 48 Uriel Jeshua Stnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) White RE (1993) A revision of the subfamily Criocerinae (Chrysomelidae) of North America north of Mexico. Technical Bulletin 1805. United States Department of Agriculture, Agri- cultural Research Service, Washington, DC, 158 pp. Wilcox JA (1965) A synopsis of the North American Galerucinae (Coleoptera: Chrysomeli- dae). New York State Museum and Science Service Bulletin 400: 1-226. Wilcox JA (1972) A review of the North American chrysomeline leaf beetles (Coleoptera: Chrysomelidae). New York Museum and Science Service Bulletin 421: 1-37. Wilcox JA (1975) Checklist of the Beetles of Canada, United States, Mexico, Central America and the West Indies. Volume 1, pt. 7. The leaf beetles (red version). Biological Research Institute of America, New York, 166 pp. Wolda H (1987) Altitude, habitat and tropical insect diversity. Biological Journal of the Lin- nean Society 30(4): 313-323. doi: 10.1111/j.1095-8312.1987.tb00305.x Wolda H, O’Brien CW, Stockwell HP (1998) Weevil diversity and seasonality in tropical Pan- ama as deduced from light-trap catches (Coleoptera: Curculionoidea). Smithsonian Con- tributions to Zoology 590: 1-79. doi: 10.5479/si.00810282.590 Appendix | Table IA. 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 1, 2, 3 and Figures 2—5) for detailed data (coordinates, elevation, type of vegetation, spatial location) of each site-plot. Month column: numbers in parenthesis refer to the total abundance obtained for that month. Marked (*) species were obtained only by collecting, independent of the standardized sweeps. NR=New record for Mexico. 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 Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... Site [plot] Site 1 [1, 2, 4, 7, 9, 11] Sited- (ly 2; 3965 7 Ose] Site 4 [3, 4, 9] Site 6 [7, 8, 11, 12] Taxon Brachycoryna pumila Guérin- Méneville, 1844 49 Month (abundance) May (2), Jun (1), Jul (1), Sep (3), Oct (2) Jan (2), Feb (2), Apr (2), Jun (1), Jul (3), Aug (1), Sep (2), Oct (2) May (1), Jul (1), Sep (1), Oct (1), Nov (1), Dec (1) Jan (1), Feb (2), May (1), Jun (2), Jul (1), Aug (4) Jul (1), Aug (1) Sep (1), Oct (1) Jan (1), Feb (1), Jul (1), Sep (1), Oct (1), Nov (2) Site 7 [1, 5] Chalepus verticalis (Chapuis, Site 7 [6] 1877) Sumitrosis inaequalis (Weber, Site 1 [6, 7, 8] 1801) 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, 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) [ [ 11, 12] [ [ Site 6 [1, 3, 9] May (1), Jul (4) Tribe Mesomphaliini Hope, 1840 Ogd j Boh : ledoecosta juvenca (Boheman Site 4 [8, 9] Jun (3), Jul (2), Aug (1) 1854) Tribe Ischyrosonychini Chapuis, 1875 Physonota alutacea Boheman, Site 1 [4] Sep (1), Oct (1) 1854 Tribe Cassidini Gyllenhal, 1813 Coptocycla (Psalidonota) texana Site (A, 2512 | (Schaeffer, 1933)* eee Helocassis clavata (Fabricius, Site 2 [5] 1798) Site 3 [4] Site 5 [3] Helocassis crucipennis (Bohe- Site 5 [7] man, 1855) Site 6 [1] he bilimeki Spaeth, Site 4 [8, 9, 10, 11] CHRYSOMELINAE Latreille, 1802 Tribe Chrysomelini Latreille, 1802 Subtribe Doryphorina Motschulsky, 1860 Jun (1), Sep (7), Oct (7), Nov (3), Dec (2) May May (1), Jul (1) May (1) Jun (1) Aug (1) Jul (1) May (1), Jul (3), Aug (3), Sep (4), Oct (3) Labidomera suturella Chevrolat, | Site 3 [3] Nov (1), Dec (1) 1844 Site 4 [11] Jul (1) Subtribe Chrysomelina Latreille, 1802 Plagiodera semivittata Stal, 1860 | .. Site 2 [5, 10, 12] Jan (1), Feb (1), Jun (2), Sep (2), Oct (2), Nov (1), Dec (1) Site.3: [3,.5;-75 8] Jan (1), Feb (1), May (2), Jun (6) Site 4 [2, 3] Apr (1), Sep (1), Oct (1) Plagiodera thymatoides Stal, Site 2 [8, 10, 11] Jun (2), Jul (1), Aug (2) 1860 Site 3 [3, 8] Jun (2) 50 Taxon GALERUCINAE Latreille, 1802 Tribe Galerucini Latreille, 1802 Group Coelomerites Chapuis, 1875 Coraia subcyanescens (Schaeffer, Site [plot] Uriel Jeshua Stnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) Month (abundance) 1906) Site 1 [1] Jun (1) Miraces aeneipennis Jacoby, Site 1 [4, 10, 11] Aug (1), Sep (2), Oct (2), Nov (2), Dec (2) 1888 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, | _. Site 7 [1, 6, 10] 1906) 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 _| Site 1 [5] & Smith, 1980 Site 3 [5] Site 4 [4] Gynandrobrotica lepida (Say, site > [L265 7, 83.110; 1835) 11,12] Site 6 [1, 3, 4, 6, 10, 112] Group Cerotomites Chapuis, 1875 Cyclotrypema furcata (Olivier, Site 4 [6, 10, 12] Sep (1), Oct (1) Sep (1), Oct (1) Aug (2) Apr (1), Aug (3), Sep (11), Oct (9), Nov (6), Dec (5) Jul (1), Aug (2), Sep (12), Oct (9), Nov (3), Dec (2) Jun (2), Aug (2) Aug (1) Sep (1), Oct (1) Jan (2), Feb (2), Nov (1) Mar (1) 1808) Site 5 [6] Tribe Alticini Newman, 1835 Acallepitrix sp. 1 Site 1 [12] Site 2 [3, 8] Site 3 [9] 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, Jan (3), Feb (8), Mar (11), Apr (1), Jul (2), Aug (2) 8, 9] 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) Jul (2), Aug (1), Sep (2), Oct (2) Jun (1), Jul (3), Aug (4), Sep (4), Oct (2) Sep (1), Oct (1) Jun (2), Sep (1), Oct (1) Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... af Taxon Site [plot] Month (abundance) Acrocyum dorsalis Jacoby, 1885 | Site 2 [3, 10] Mar (1), Sep (1), Oct (1) Site 3 [2] Jan (1), Feb (1) sis Sein a cna aaoudina |S | Apr (1), Sep (1); Oct (1) ricius, 1801) 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, | Site 1 [2, 3, 6, 12] Jan (2), Feb (3), Jun (2), Aug (1), Sep (1), Oct (1) 1877) 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, 2s 3, 4, 5, 6, 7.,- B50 NOs Ts. 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, a5 6, 7s 8, Jan (8), Feb (17), Mar (3), Apr (1), May (2), Jun 9, 10, 11, 12] (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. | 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) 52 Chrysogramma sp. 1 Taxon Site [plot] Site 1 [6] Site.2 [1, 2, 3, 4, 6, 7, 951.0, 11 2] Site’3 [1, 2; 3, 4, 5,6, 79.85 10; 1 yl] Uriel Jeshua Stnchez-Reyes et al. / ZooKeys 611: 11-56 (2016) Month (abundance) Apr (1) Jan (12), Feb (41), Mar (45), Apr (6), May (2), Jul (1) Jan (14), Feb (34), Mar (26), Apr (8), May (3) Dibolia sp. 1 Site 4 [4] Aug (1) lie i glabrata (Fabricius, Site 5 [1, 6] Aug (2) Disonycha stenosticha Schaeffer, | Site 1 [6] Nov (2) ree a ANB, ASB Mo |), Sep (2), Ob @): Nov 5) Dee (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) Site74 (25-9 | 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, | Jan (2), Feb (2), Mar (1), May (1), Jun (2), Jul (1), 8, 12] 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) ‘3 PUN Be ROTI ae ay abv) Mans Jul Ys Ants (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, | Jan (3), Feb (4), Mar (5), Apr (1), May (1), Jun (1), 9, 10] Site.5 [15 355;'3] Site 6 [5, 10] Site 7 [1] Jul (5), Aug (2), Sep (3), Oct (1) May (1), Aug (3) Jul (2), Aug (1) Jul (1) Faunistic patterns of leaf beetles (Coleoptera, Chrysomelidae) within elevational... 53 Taxon Site [plot] Month (abundance) Lupraea sp. 1 Site 3 [5] Sep (1), Oct (1) - a 4557-8 | 45.03), 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 on Mar (1) Margaridisa atriventris Site2 (2059 LI Aug (3) (Melsheimer, 1847) Site 3 [1, a 6, 8, 10, | Jan (1), Feb (1), May (1), Jun (1), Jul (7), Aug (14), 1d, 12] 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, Site 3 [3] Aug (1) 1891 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) fe a, 2,3, 4, 5, Apr (7), May (6), Jun (7) Sela DaTS SUES