San Buenaventura (from here on SBV) is located on the eastern slope of the Sierra de Cacoma-Sierra de Manantlan, Jalisco, Mexico (latitude 19°45'19'', 19°48'50'' N and longitude -104°01'25'', -104°08'25'' W; Fig. 1). The climate is warm sub-humid type according to the Köeppen climate classification modified by García (1988). The average annual precipitation from the nearest weather station Presa Basilio Vadillo (Comisión Nacional del Agua and Servicio Metereológico Nacional 2023) was 747 mm, while the average air temperature was 23.8°C, with average maximum and minimum temperatures of 31.9°C and 15.8°C respectively. The highest temperature during the study period was recorded in May and the lowest in January (Fig. 2). The dominant vegetation in the area is TDF. The dominant tree species are Lysiloma acapulcense (Kunth) Benth (Fabaceae), L. divaricatum (Jacq.) J.F.Macbr, Jacaratia mexicana A. DC. (Caricaceae), Amphipterygium adstringens (Schltdl.) Standl. (Anacardiaceae), Entada polystachya (L.) DC. (Fabaceae), Ceiba aesculifolia (Kunth) Britten & Baker f. (Malvaceae), Senegalia macilenta (Rose) Britton & Rose (Fabaceae), Vitex mollis Kunth (Lamiaceae), Ipomoea bracteata Cav. (Convolvulaceae), Bursera spp. (Burseraceae) and Cochlospermum vitifolium (Willd.) Spreng. (Bixaceae) (Jardel 1992). A gallery forest, characterised by trees taller than those of the TDF, extends along streams and narrow canyons. Flat areas in the zone have been open to agriculture and hillsides are used as grazing areas for cows and goats, which has resulted in the near disappearance of the native understorey.

Figure 1.  

Sampling localities of Odonata species in San Buenaventura, Jalisco, Mexico. Entomological samplings were performed from 1996-1997. Imagery 2015, INEGI Maxar Technologies CNES/Airbus. Downloaded August 2023.

Figure 2.  

Monthly variation of rainfall and temperature in San Buenaventura, Jalisco during 1996-1997.

The study area is located within the Ayuquila-Armeria River Basin and, more specifically within the Tuxcacuesco River and Ayuquila sub-basins (Guevara Gutiérrez et al. 2019, Rodríguez-Contreras et al. 2019). Sampling collections were done around three main localities: SBV town, Los Yesos and Amacuahutitlán, and only occasionally at Las Higueras (see Table 1, Fig. 1).

The SBV samplings were made along the margins of the Ferreria River, a permanent river that crosses the town. Samplings in Amacuahutitlán and Los Yesos were done at small open streams and finally, at the site of Las Higueras consisting of a small, mostly shaded, spring-fed shallow stream with abundant aquatic plants on its surface (Fig. 3). Most collecting sites belong to the Municipality of El Limón, except for Amacuahutitlán, which is located within the Municipality of Tonaya (see Table 1).

Figure 3.  

Habitats and Odonata species from four localities in San Buenaventura, Jalisco, Mexico; a, b microhabitat of Anisagrion allopterum; c Dythemis maya; d Neoneura amelia; e Gynacantha helenga; f Archilestes grandis; g Progomphus clendoni. Photos: a, b, César Durán; c, f, g Enrique González Soriano; d, Enrique Ramírez; e, Eric Hough (Naturalista).

Fieldwork in SBV was always conducted by two people between November 1996 and October 1997. Collections were carried out for a period of five days every month. Specimens were obtained through direct collecting, between 09:00 h and 15:00 h (10:00 h-16:00 h in the daylight-saving time).

All the Odonata records collected in SBV, Jalisco, Mexico for this work were included in the GBIF dataset Digitization and Systematization of the National Biological Collections of the Institute of Biology, UNAM from the National Autonomous University of Mexico (Sánchez Cordero Dávila 2023). Data on the phenology and the number of specimens collected (between parentheses) were added in the Odonata species list (Results section); additional information can be consulted in the Supplementary materials and photographs of some species in Fig. 3.

The diversity of the SBV odonate assemblage was analysed through different metrics of species and phylogenetic diversities. We first quantified:

(a) abundance, measured as the number of specimens collected through a species rank abundance curve;

(b) species richness, as the number of species observed (diversity order 0, ⁰D);

(c) Shannon diversity, which corresponds to the exponential of the Shannon Index (diversity order 1, ¹D); and

(d) Simpson diversity, which corresponds to the inverse of the Simpson Index (diversity order 2, ²D) (Jost 2006, Chao et al. 2014).

The measurement unit for ¹D and ²D is the number of effective species, also referred to as Hill numbers, in such a way that ¹D indicates the effective number of equally abundant species within an assemblage and ²D showed the effective number of the most abundant or most dominant equally abundant species.

We then calculated the maximum expected richness value of diversity for ⁰D, ¹D and ²D to compare those values with our observed sample. We used the Spade R package (Chao et al. 2015) to compute the non-parametric abundance-based Chao 1-bias corrected estimator and the estimators proposed by Magurran for ⁰D, ¹D and ²D, respectively (Magurran 1988, Chao 2005, Chao et al. 2013). We also calculated a cumulative species curve for the whole Odonata assemblage from SBV, through the interpolation-extrapolation method proposed by Chao et al. (2014) for ⁰D, ¹D and ²D and using the iNEXT R package (Hsieh et al. 2016).

In addition, we evaluated monthly abundance, ⁰D, ¹D, and ²D to analyse temporal diversity patterns in the Odonata assemblage. In addition, we analysed temporal phylogenetic diversity through the taxonomic diversity (Δ) and taxonomic distinctness (Δ*) indices, which are based on the abundance and the average taxonomic distinctness (Δ+) index, based on species incidence (Warwick and Clarke 1995, Clarke and Warwick 2001). Taxonomic diversity and taxonomic distinctness analyse phylogenetic divergence amongst species within communities or assemblages according to their topological organisation, i.e. the phylogenetic relationships amongst taxa and their taxonomic hierarchy; these measures calculate how closely related the specimens (Δ) or the species within the assemblage are (Δ*) or how evenly distributed their evolutionary paths are through the taxonomic hierarchy (Δ+) (Clarke and Warwick 1998, Clarke and Warwick 2001). Since the most recent odonate phylogenies do not include all taxa within the order (e.g. Bybee et al. (2021)), we used the hierarchical classification above species-level as a proxy to calculate taxonomic distances amongst taxa using diferent phylogenetic diversity metrics (Clarke and Warwick 2001; Tucker et al. 2016). In particular, the phylogenetic hypothesis of Bybee et al. (2021) for suprafamily levels and Dijkstra et al. (2014) and Carle et al. (2015) for suprageneric levels of Anisoptera and Zygoptera suborders, respectively were used for this purpose. We included seven taxonomic levels to evaluate monthly phylogenetic divergence within the SBV odonate assemblage: order, suborder, superfamily, subfamily, tribe, genus and species. Phylogenetic divergence was calculated through the taxondive and taxa2dist functions of the Vegan R package (Oksanen et al. 2015).

To analyse whether species and phylogenetic diversity of the SBV odonate assemblage are related to abiotic factors, we performed Pearson’s correlation analyses between monthly species diversity (abundance, ⁰D, ¹D and ²D), monthly phylogenetic diversity (Δ, Δ*, Δ+) and monthly mean rainfall and temperature documented in SBV, Jalisco during the sampling time. Values of precipitation and temperature were obtained from the closest weather station (Presa Basilio Vadillo) through the National Meteorological System (Comisión Nacional del Agua and Servicio Metereológico Nacional 2023). We found a positive correlation between temperature and precipitation. Pearson’s correlation analyses were done in Past software (Hammer et al. 2001). In addition, we generated a heatmap using the pheatmap R package (Warnes et al. 2012) to display the presence and abundance that each odonate species showed monthly. Monthly diversity and phylogenetic divergence analyses allowed us to evaluate how the species diversity and the taxonomic assemblage structure were related to monthly changes of temperature and humidity. In other words, those analyses allowed us to evaluate how the abiotic factors can be associated with the temporal structure of the Odonata community of the TDF.

We documented a total of 1087 specimens belonging to seven families, 35 genera and 66 species of odonates in the assemblage (Figs 4, 5). Those values represent 87.5% of the families, 80% of the genera and 51% of the total species previously reported for the State of Jalisco (González-Soriano and Novelo-Gutiérrez 2014; Suppl. material 1). Libellulidae and Coenagrionidae were the families with the highest number of species, with 28 (42.4%) and 21 (31.8%), respectively, followed by Gomphidae (7), Aeshnidae (6), Calopterygidae (2), Lestidae (1) and Platystictidae (1). At the generic level, Libellulidae had the highest numbers of genera (15), followed by Coenagrionidae (8), Gomphidae (5), Aeshnidae (4), Lestidae (1), Calopterygidae (1) and Platystictidae (1). Argia was the most speciose genus with 10 species, followed by Macrothemis with four and the remaining genera with 1-3 species. Phyllogomphoides luisi was recorded for the first time in the State of Jalisco and Anisagrion allopterum represented the first northernmost documented record of this species in America (Fig. 3). The total species richness (66 species) represented 88.9% from the total expected richness (⁰D, 74.16 effective species), 96.3% (35.4) of the expected Shannon diversity (¹D) and 97.97% (22.2) of the expected Simpson diversity (²D) (Table 2, Fig. 5). The estimated species richness suggests that there could be another eight species in the site, whereas observed evenness (¹D) and dominance (²D) showed values close to the estimated values of those diversity metrics.

Figure 4.  

Species rank-abundance curve for the odonate species collected in San Buenaventura, Jalisco during 1996-1997.

Figure 5.  

Interpolation-extrapolation cumulative curve, based on the number of odonate specimens collected in San Buenaventura, Jalisco (1996-1997) for three diversity orders: ⁰D, species richness; ¹D, Shannon diversity; ²D, Simpson diversity. Exp, expected values of diversity of order 0, 1 and 2.

Species abundance during all the sampling was very heterogeneous. Only a few species were very abundant and most were represented by one or few specimens (Figs 4, 6). Hetaerina americana was by far the most abundant species with 149 specimens, followed by the coenagrionids Argia pulla (68), A. extranea (59), Phyllogomphoides pacificus (51), Enallagma novaehispaniae (49) and A. harknessi (44). Those six species represented 41% of the total abundance of the assemblage (420 specimens). Surprisingly, one anisopteran, P. pacificus, appeared within the group of abundant species with more than 50 specimens. This contrasts with our other previous studies in TDF where the most abundant species belong to the suborder Zygoptera. On the contrary, 16 species were represented by only 1-3 specimens and contributed only 3% of the total abundance (31 specimens).

Figure 6.  

Heatmap showing monthly abundance (number of specimens) of odonate species collected in SBV, Jalisco. Species in rows are ordered according to their suborder and family. AES, Aeshnidae; GOM, Gomphidae; LIB, Libellulidae; CAL, Calopterygidae; COE, Coenagrionidae; LES, Lestidae; PLA, Platystictidae.

We observed a high variation in the different facets of diversity species of the odonate assemblage throughout the year. The highest species richness was recorded in August (44 species) and September (40), while the lowest was observed in November (11) and January (11). In addition, the highest value of abundance was observed in June (202 specimens), during the rainy season and the lowest in November (17) and May (21), during the dry season (Table 2, Fig. 6). The highest Shannon diversity was also documented in August and September, while the highest value of Simpson diversity was recorded in September (Table 2).

Phylogenetic divergence also showed a high variation throughout the year (Table 2): the taxonomic diversity showed its highest values from July to October; the taxonomic distinctness (Δ*) was higher in June and October. In general, it was high throughout the rainy period, whereas we observed lower values than the expected (76.42 species) from January to April except December. In general, the monthly taxonomic diversity (Δ) and the average taxonomic distinctness (Δ+) were lower than the expected.

Variation in precipitation showed a positive, moderate correlation with Shannon and Simpson diversities of the SBV odonate assemblage, as well as with phylogenetic divergence (Δ* and Δ+) (Table 3); whereas variation in minimum temperature also showed a positive, moderate correlation with monthly variation in species richness, Shannon diversity and all metrics of phylogenetic diversity (Table 3). Maximum temperature did not show any influence on the different metrics performed.

Monthly precipitation was strongly correlated with monthly minimum temperature (r = 0.833, P < 0.001); abundance showed a high, positive correlation with species richness and Shannon diversity (Table 3), which, in turn, were strongly correlated with Simpson diversity; also, taxonomic diversity and taxonomic distinctness showed a high, positive correlation between them. In addition, average taxonomic distinctness was highly and positively correlated with the diversity metrics of most species.

In contrast with other TDF Mexican localities studied where coenagrionids were dominant (e.g. Dominguillo, Oaxaca; Huautla, Morelos; San Javier, Sonora), the odonate assemblage from SBV was dominated by one abundant calopterygid species: Hetaerina americana (Fig. 2). In addition, odonate species richness in SBV (66 species) was higher than the richness reported from Sierra de Huautla, Morelos (57) (González-Soriano et al. 2008); San Javier, Sonora (52) (González-Soriano et al. 2009); Río Pinolapa, Michoacán (51) (Novelo-Gutiérrez and Gómez-Anaya 2009); Dominguillo, Oaxaca (50) (González-Soriano et al. 2021); and Aguililla, Michoacán (40) (Novelo-Gutiérrez and Gómez-Anaya 2009). However, it was lower than that reported for Chamela, Jalisco (González-Soriano et al. 2004), which is the TDF site with the largest number of species recorded so far (78). SBV shares 71.2% species with Huautla; 63.6% with Chamela; 56.6% with Río Pinolapa; 53% with San Javier; 48.5% with Aguililla; and 42.4% with Dominguillo.