Neuroptera Diversity from Tacan á Volcano, Mexico: Species Composition, Altitudinal and Biogeographic Pattern of the Fauna

: Approximately 340 species of ten families of Neuroptera have been recorded from Mexico. The Tacan á volcano, reaching an elevation of 4092 m a.s.l., is the northernmost representative of the Central American Nucleus volcanic range. Recent survey efforts of the Neuroptera diversity of the Tacan á volcano, Chiapas, along an altitudinal gradient, increased the known fauna of this order in Mexico by 31 species and two genera: Biramus Oswald, 1993 (Hemerobiidae), and Titanochrysa Sosa & Freitas, 2012 (Chrysopidae), with extension of the known distributional range of 25 species of ﬁve families. Most of the new country records are from species previously known only from Central and South America. The lacewing fauna of Chiapas is updated from 91 to 147 species. The Neuroptera of the Tacan á volcano is mostly Neotropical with some taxa of Nearctic afﬁnity restricted to medium and high elevations. More than 80% of the Tacan á volcano lacewing species also occur in the Brazilian subregion, especially the Mesoamerican and Paciﬁc domains. Neuropteran species were recorded from 650 to more than 3500 m a.s.l. A higher species richness was present between 600 and 1700 m, with a few species occurring at altitudes above 3000 m. A species checklist and an identiﬁcation key to the genera of Neuroptera of the Tacan á volcano are provided.

Studies on the Neuroptera fauna of Mexico have been scattered for decades, with infrequent works by European and American entomologists that collaterally built a record of the fauna. Notably, the Spanish Jesuit priest Longinos Navás described several species between 1911 and 1936, followed by the works of other American, European, and Latin American entomologists (Table 1).

Sampling
Sampling was carried out monthly between February 2018 and January 2019, at different sites and altitudinal levels with different types of vegetation (evergreen tropical forest, coffee plantation, cloud forest, oak forest, and pine forest). Specimens were captured at each collecting station using a black and mercury vapor light trap (screen) and bucket (black light), two Malaise traps, five ground-level interception traps, five yellow plate traps at the tree canopy, and entomological net on vegetation ( Figure 1). Sporadic sampling also was applied using light traps and entomological nets. Specimens were kept alive in plastic screw cap vials, transported to the laboratory, and then pinned or conserved in 80% ethyl alcohol.  1 Figure 3. Map with the sampling sites at the Tacaná volcano Biosphere Reserve (Chiapas, Mexico). A1-A5: Annual systematic sampling sites; S1-S5: Sporadic sampling sites. Design of the wind rose was based on the Mayan symbol "the Four Sides of the Earth (Xocom Balumil)"; the rhombuses on the vertical axis joined to the central rhombus mean east and west, while the extreme lateral rhombuses represent the north and south [redrawn from [152].

Parsimony Analysis of Endemism (PAE)
To assist in a better understanding of the altitudinal distribution of the Neuroptera species from the Tacaná volcano, we performed different PAE analyses. PAE constructs cladograms based on the cladistic analysis of the presence-absence data matrices of species and supraspecific taxa [153]. In this analysis, a matrix was built with distributional units used as the "terminals", and the taxa (species, genus, family, etc.) used as "characters", so a parsimony analysis is performed, resulting in the most parsimonious cladograms used to describe a potential pattern of relationship of the distributional units (e.g., areas of endemism, altitudinal levels, etc.). Initially, we performed a PAE using the sites at the main five levels of sampling as terminals, and the Neuroptera species present as characters; additional PAE were carried out for each Neuroptera family in order to unravel possible altitudinal influences from each group; the exception was Rhachiberothidae, which has only a representative species in this study.
Species ("characters") were codified as present (1) or absent (0) for each of the distributional units ("terminals"). A hypothetical distributional unit with absence of all species was used for rooting the tree. The matrices (Appendix A) were built with WinClada [154], and then exported as a Nexus file to perform phylogenetic analysis under the principle of parsimony in TNT (Tree Analysis using New Technology, version 1.5) [155]. The most parsimonious cladogram was obtained through heuristic algorithms using the tree bisection and reconnection method (TBR), using as parameters the following: random seed = 0, hold = 3000, and hold/ = 50 of 60 replications. The most parsimonious topology (or the strict consensus of the most parsimonious topologies) was exported to Illustrator CS6 software to be edited.

Distribution Map
Design of the study site map and the location of the sampling points was done using ArcGIS 10.2.2. The different layers (federative entities and municipalities) used for this map were obtained from the information provided by the Instituto Nacional de Estadistica y Geografia (INEGI), Mexico. Statistical and geographical information is at a scale of 1:50,000. The projection of localities with geographical coordinates was carried out with UTM (Universal Transverse Mercator). Subsequently, a raster of the model of the Mexican Continuum of Elevations 3.0 [157] of the area of Chiapas was built, with a cut of the municipalities that were within the study area, using a layer of vectors of the municipal boundaries of the state. After the area of study was selected, adjustments were made to the elevation model with a reclassification of values of z (altitude) to be able to visualize the altitude difference within the area of interest. Within the reclassification, seven intervals ranging from 0 m to 4080 m were used. In addition, a shadow map (hillshade) was made that helped us to better visualize the slopes of the terrain where the study area was selected.

Results
A total of 2534 specimens from 109 species of 28 genera belonging to six families of Neuroptera were collected. Thirty-one species and two genera of Neuroptera, Biramus Oswald, 1993 (Hemerobiidae), and Titanochrysa Sosa & Freitas, 2012 (Chrysopidae), were recorded for the first time from Mexico; 25 species were recorded for the first time from Chiapas state. Such data increase the Mexican lacewing fauna from 343 species of 77 genera to 374 species of 79 genera belonging to 10 families. Chrysopidae, Coniopterygidae, and Hemerobiidae present most of the new records. Currently, the known fauna of Neuroptera

Meleoma titschacki Navás, 1928
Distribution: Costa Rica and Mexico (Chiapas) [11,14,80]. Remarks: Meleoma titschacki presents a Neotropical distribution. Specimens were from cloud forest and oak forest, with some specimens found on Quercus spp. and Saurauia spp. Flight period was from January to December (in both seasons). Previous altitudinal distribution records were between 640 and 1990 m [80]; herein, altitudinal records were from 1222 to 2142 m, the highest altitudinal distribution recorded for this species.

Genus
Remarks: Plesiochrysa brasiliensis has a broad Nearctic and Neotropical distribution. Specimens were collected in evergreen tropical forest, cloud forest, and mixed oak-cloud forest. Flight period was in January to March, May to July, October, and November (in both seasons). Previous altitudinal distribution records are between sea level and 1800 m [80]; herein, specimens were collected from 661 to 2060 m, the highest altitudinal record for P. brasiliensis.

Plesiochrysa sp. 1
Distribution: Mexico (Chiapas). Remarks: Specimens were collected in cloud forest and oak forest, with flight period in May, July, and October (in the rainy season), and an altitudinal record at 2081 m. These female specimens present a longer pronotum and spermatheca different from P. brasiliensis and P. sp. 2, being more flattened at the base and with wider vela.

Plesiochrysa sp. 2
Distribution: Mexico (Chiapas). Remarks: Specimens were collected in evergreen tropical forest and cloud forest, with flight period from January to March (in the dry season), and altitudinal records from 661 to 1217 m. These female specimens are similar to Plesiochrysa sp. 2 of Sosa [208], a species not yet formally described by this author.

Genus Titanochrysa Sosa & de Freitas, 2012
Titanochrysa annotaria (Banks, 1945) Distribution: Costa Rica, Panama [11,139,142,158,201], and Mexico (new record) (Chiapas). Remarks: Titanochrysa annotaria was previously recorded from Costa Rica and Panama. This is the northernmost record for the species, ca. 1178 km north from previous records (La Amistad International Park, Puntarenas, Costa Rica). This species, together with T. simpliciala Tauber et al., 2012a, confirm the first occurrence of the genus Titanochrysa in Mexico. Specimens were collected in cloud forest, with some specimens found on Citrus spp. and Saurauia spp. Flight period was in January and June to November (mainly in the rainy season). Previous altitudinal distribution records are between 1500 and 1600 m [201]; herein, specimens were collected from 1250 to 1577 m, which is the lowest altitudinal record. Variation: Individuals usually have the small cluster of veinlets in the middle of the inner gradate series of the forewing evident, but some studied specimens had this cluster of veinlets reduced or not very evident.

Titanochrysa simpliciala Tauber et al., 2012
Distribution: Costa Rica [11,139,142] and México (new record) (Chiapas). Remarks: Titanochrysa simpliciala was previously recorded from Costa Rica. This is the northernmost record for the species, ca. 1032 km north from previous records (Quebrada Amistad, Heredia, Costa Rica). Specimens were collected in cloud forest, with some specimens found on Miconia spp. Flight period was in March and November (mainly in the dry season). A previous altitudinal distribution record is at 1920 m [139]; herein, specimens were collected from 1565 to 1625 m, which are the lowest altitudinal distribution records for this species.  [11,27,80,158,209], and Mexico (new record) (Chiapas).

Genus
Remarks: Leucochrysa (L.) clara has a Neotropical distribution. This is the northernmost record for the species. Specimens were collected in cloud forest. Flight period was in January to February, May, and August to October (mainly in the rainy season). Previous altitudinal distribution records are from lowlands (7 to 458 m) [27,80]; herein, altitudinal distribution was from 1080 to 1705 m, which is the highest altitudinal distribution record.
Remarks: Leucochrysa (L.) colombia presents a Neotropical and Nearctic distribution, with several Neotropical records. Herein, we record this species for the first time in Mexico, in the state of Chiapas. Specimens were collected in cloud forest and oak forest. Flight period was in April and May (mainly in the dry season). Previous altitudinal distribution records are between 1500 and 2700 m [80,190]; herein, specimens were collected between 1705 and 2079 m.

Leucochrysa (Leucochrysa) lestagei Navás, 1922
Distribution: Brazil, Costa Rica, Ecuador [11,80,210], and Mexico (new record) (Chiapas). Remarks: Leucochrysa (L.) lestagei presents a Neotropical distribution. This is the northernmost record for the species, ca. 1182 km north from the previous record (La Amistad International Park, Puntarenas, Costa Rica). The studied specimen was collected in cloud forest, in June (in the rainy season). Previous altitudinal distribution records are from mid-elevations and lowlands [80]; herein, the altitudinal record of this species was 1440 m.
Remarks: Leucochrysa (L.) pretiosa presents a Neotropical distribution. Specimens were collected in evergreen tropical forest and coffee plantations, with some specimens found on Coffea sp. and Inga sp. Flight period was in January and July to December (mainly in the rainy season). Previous altitudinal distribution records are between sea level and 1300 m [80,190,211]; herein, specimens were collected between 661 to 809 m.
Remarks: Tauber et al. [141] confirmed the presence of L. (L.) varia only in South America. Nevertheless, a single male specimen, collected in cloud forest, confirms the presence of this species in Mexico for the first time. This record expands the species range ca. 2460.55 km north from records of Tauber et al. [141]. Flight time was in July (in the rainy season) and the altitudinal record was 1254 m, higher than previous records (200 to 916 m) [141]. Leucochrysa (L.) varia was previously recorded in Mexico (Tabasco) by Navás [27] and Adams [28] also mentioned its probable occurrence in Mexico. However, Tauber et al. [141] sustained that such previous records are probably misidentifications. Herein, we confirm the presence of the species in southern Mexico.

Leucochrysa (Leucochrysa) variata (Navás, 1913)
Distribution: Mexico (Chiapas (new record), Veracruz) and Panama [11,27,141]. Remarks: Leucochrysa (L.) variata presents a Neotropical distribution. Specimens were collected in evergreen tropical forest, coffee plantations, and cloud forest, with some specimens collected on Citrus spp. Flight period was in January, October, and November (mainly in the dry season). Previous altitudinal distribution records are between 7 to 122 m [27]; herein, specimens were collected from 789 to 1585 m, which is the highest altitudinal record for this species.

Leucochrysa (Nodita) amistadensis Penny, 2001
Distribution: Costa Rica [11,80] and Mexico (new record) (Chiapas). Remarks: Leucochrysa (N.) amistadensis was previously known from Costa Rica. This is the northernmost record, ca. 1220 km north from previous records (La Amistad International Park, Puntarenas, Costa Rica). Specimens were collected in cloud forest and mixed oak-cloud forest. Flight period was in May, June, and September (mainly in the rainy season). Previous altitudinal distribution records are between 1500 and 1600 m [80,159]; herein, specimens were collected from 1440 to 2120 m, which is the highest altitudinal record for this species.
Remarks: Leucochrysa (N.) askanes presents a Neotropical distribution. Specimens were collected in evergreen tropical forest, coffee plantations, and cloud forest, with some specimens collected on Inga spp. and Coffea spp. Flight period was in January and April to December (mainly in the rainy season). Previous altitudinal distribution records are between 40 and 1500 m [80,158]; herein, specimens were collected from 680 to 1085 m.

Leucochrysa (Nodita) azevedoi Navás, 1913
Distribution: Brazil [11,29,166,204] and Mexico (new record) (Chiapas). Remarks: Leucochrysa (N.) azevedoi presents a Neotropical distribution. This is the northernmost record for the species, ca. 6885 km north of previous records (Est. Exp. PESAGRO, Campos dos Goytacazes, Rio de Janeiro, Brazil). A single male specimen was collected in evergreen tropical forest. Its flight date was in January (in the dry season) and its altitudinal record was 743 m, higher than the previous record for the species [29].

Leucochrysa (Nodita) camposi (Navás, 1933)
Distribution: Ecuador [11,29,214] and Mexico (new record) (Chiapas). Remarks: Leucochrysa (N.) camposi was previously known from Ecuador. This is the northernmost record for the species, ca. 2343 km north from previous records (Guayaquil, Ecuador). Specimens were collected in cloud forest. Flight period was in June and December (in both seasons) and their altitudinal distribution records were from 1231 to 1620 m.
Remarks: Leucochrysa (N.) caucella presents a Neotropical distribution. Herein, we record this species for the first time in Mexico. This is the northernmost record for the species, ca. 1220 km north from previous records (La Amistad International Park, Puntarenas, Costa Rica). Specimens were collected in cloud forest, with some specimens collected on Quercus spp. Flight period was in February and October (in both seasons). Previous altitudinal distribution records are from 914 to 1982 m [21,80]; herein, specimens were collected from 1557 to 1582 m. Variation: The original description of L. (N.) caucella does not include characteristics of terminalia and genitalia, and mentions that the wing-mesothorax connection lacks a dark spot, but some specimens had dark spots on this area, as well as on the ectoprocts.

Leucochrysa (Nodita) digitiformis Tauber et al., 2008
Distribution: Brazil [11,138] and Mexico (new record) (Chiapas). Remarks: Leucochrysa (N.) digitiformis was previously known from Brazil. This is the northernmost record for the species, ca. 6534 km north from previous records (Campos dos Guytacazes, Rio Grande do Sul, Brazil). Specimens studied were from coffee plantations and cloud forest. Flight time was in July (in the rainy season). Previous altitudinal distribution records are between 14 and 30 m [138]; herein, specimens were collected at 720 m, which is the highest altitudinal record for this species.

Leucochrysa (Nodita) lateralis Navás, 1913
Distribution: Brazil, Guatemala [11,27,137] and Mexico (new record) (Chiapas). Remarks: Leucochrysa (N.) lateralis was previously known from Guatemala and Brazil. Herein, we record this species for the first time in Mexico, in the state of Chiapas. This is the northernmost record for the species, ca. 119.34 km north from previous records (Atitlán Volcano, Guatemala). Specimens were collected in evergreen tropical forest and coffee plantations, with some specimens found on Coffea spp. Flight period was in January, July, and October to December (mainly in the dry season). Previous altitudinal distribution records are between 7 and 1067 m [27]; herein, altitudinal distribution records were from 713 to 780 m. Variation: Specimens of this species typically have red gena, frons with short red bands below the antennal base, a completely dark dorsal surface of the scape, and a Vor Y-shaped dark mark behind the antenna on the vertex, but some studied specimens had pale gena, a scape with red spots, lack of marks on the frons, and reduced Y-or V-shaped marking on the vertex.

Leucochrysa (Nodita) maculosa de Freitas & Penny, 2001
Distribution: Brazil [11,137] and Mexico (new record) (Chiapas). Remarks: L. (N.) maculosa was previously known from Brazil. This record is the northernmost record for the species, ca. 6247 km north from previous records (Taquaritinga, São Paulo, Brazil). Specimens were collected in evergreen tropical forest and cloud forest. Flight period was from April to July, September, and December (mainly in the dry season). Previous altitudinal records are unknown. Specimens were collected from 753 to 1736 m. Variation: Individuals of this species typically have a pale maxillary palp basally, dark on the fourth and basal half of the apical segment, pale on the apical half, a pale clypeus, wine red marks below the antennal base from the eye margin, a mesonotum with a brick red mark along the lateral margin, and a red spot on the second tergite, but some studied specimens had a pale palp, clypeus with reduced marks, red spots in front of the antenna, without markings on mesonotum, and with a dark spot on 6th and 7th tergites.
Remarks: Leucochrysa (N.) nigrovaria presents a Neotropical distribution. Specimens were collected in evergreen tropical forest and cloud forest, with some specimens collected on Citrus spp. Flight period was in January, April, July, September, and November (mainly in the dry season). Previous altitudinal distribution records are between 457 and 1500 m [21,80,171]; herein, specimens were collected from 678 to 1250 m. Variation: Individuals of this species typically have a pale green pronotum with a longitudinal lateral reddish-brown stripe, a meso-and metanotum mottled brown and green to completely dark brown, and large reddish-brown spots on tergites 4, 6, 7, and 8, but some studied specimens had lateral pronotum stripes thicker and darker, a meso-and metanotum more or less black pigmented, and an abdomen with large black spots throughout the tergites.

Leucochrysa (Nodita) squamisetosa de Freitas & Penny, 2001
Distribution: Brazil [11,137] and Mexico (new record) (Chiapas). Remarks: Leucochrysa (N.) squamisetosa was previously known from Brazil. This is the northernmost record for the species, ca. 6095 km north from previous records (Birigui, São Paulo, Brazil). A single female specimen was collected in evergreen tropical forest. Its flight date was in June (in the rainy season) and its altitudinal record was 659 m. Previous altitudinal distribution of this species is unknown.

Leucochrysa (Nodita) tarini (Navás, 1924)
Distribution: Cuba [11,136,174] and Mexico (new record) (Chiapas). Remarks: Leucochrysa (N.) tarini was previously known from Cuba. Specimens were collected in evergreen tropical forest, coffee plantations, and cloud forest, with some specimens collected on Inga spp. and Coffea spp. Flight period was in January to March, May to July, and October (mainly in the rainy season). Previous altitudinal distribution of this species is unknown, herein its altitudinal records were from 661 to 1223 m.

Leucochrysa (Nodita) sp. 1
Distribution: Mexico (Chiapas). Remarks: Specimens of Leucochrysa sp. 1 were collected in evergreen tropical forest, coffee plantations, and cloud forest; their flight period was in January, April, July, September, and October (mainly in the rainy season), and their altitudinal distribution records were from 741 to 1483 m. They are morphologically close to Leucochrysa (Nodita) zayasi Alayo, 1968 but with evident differences in genitalia.

Leucochrysa (Nodita) sp. 2
Distribution: Mexico (Chiapas). Remarks: A single female specimen of Leucochrysa sp. 2 was collected in mixed oakcloud forest, with flight time in September (in the rainy season), and an altitudinal record of 2149 m. The specimen is morphologically close to Leucochrysa (Nodita) compar Alayo, 1968, but with evident differences in spermatheca.

Leucochrysa (Nodita) sp. 3
Distribution: Mexico (Chiapas). Remarks: A single female specimen of Leucochrysa sp. 3 was collected in evergreen tropical forest, with flight time in November (in dry season), and an altitudinal record of 713 m. The specimen is morphologically close to L. (Nodita) azevedoi Navás, 1913, but with evident differences in spermatheca.

Leucochrysa (Nodita) sp. 4
Distribution: Mexico (Chiapas). Remarks: A single male specimen of Leucochrysa sp. 4 was collected in cloud forest, with flight date in February (in the dry season), and an altitudinal distribution record of 1195 m. The specimen is morphologically close to Leucochrysa (Nodita) cerverai  but with evident differences in genitalia.

Leucochrysa (Nodita) sp. 5
Distribution: Mexico (Chiapas). Remarks: A single male specimen of Leucochrysa sp. 5 was collected in cloud forest, with flight time in April (in the dry season), and an altitudinal distribution record of 1192 m. The specimen presents genitalia similar to Leucochrysa (Nodita) incognita de Freitas & Penny, 2001, but with reduced gonocorns, and with pale palp and gena, a vertex with a reddish inverted U-shaped marking and thorax with dark markings on the meso-and metathorax.

Leucochrysa (Nodita) sp. 6
Distribution: Mexico (Chiapas). Remarks: Specimens of Leucochrysa sp. 6 were collected in cloud forest, with the flight period in September and October (in the rainy season), and an altitudinal distribution record of 1479 m. These specimens present pale antenna and gena, frons with a reddish spot between the antennae, a vertex with two wine red lines in a V-shape, the pro-, meso-, and metanotum yellowish-green with two red spots on each side, a red mark on the anterior part of mesoscutum, and the posterior part of the meso-and metanotum reddish-orange, abdomen with reddish marks dorsally, also with particular characteristics in the genitalia. Distribution: Costa Rica, Guatemala [11,16], and Mexico (new record) (Chiapas). Remarks: Neoconis dentata was previously known from Costa Rica and Guatemala. This is the northernmost record for the species. Specimens were collected in evergreen tropical forest, coffee plantations, cloud forest, oak forest, and pine forest, with some specimens collected on Saurauia spp. Flight period was in January and March to December (in both seasons). The previous altitudinal distribution of this species is unknown; herein, it was recorded from 693 to 3089 m. Distribution: Mexico (Chiapas (new record), Tlaxcala) and United States of America [9,11,16,32,143,215].

Subfamily
Remarks: Coniopteryx (S.) latipalpis presents a Nearctic distribution, with many North American records. This is the first record of the species for the Neotropical region, as well as its southernmost record, ca. 843 km south from previous records (Nanacamilpa, Tlaxcala, Mexico). Specimens were collected in mixed oak-cloud forest and pine forest, with some specimens collected on Fuchsia spp. and Pinus spp. Flight period was in February, April to July, and October (mainly in the rainy season). Previous altitudinal distribution records are between 336 and 3048 m [9,16,143]; herein, specimens were collected from 2079 to 3277 m, which is the highest altitudinal distribution record for the species. A wide distribution in Mexico is corroborated for this species, with an affinity for high-altitude and pine forest vegetation.
Remarks: Coniopteryx (C.) simplicior presents a Nearctic and Neotropical distribution. Herein, we record this species for the first time in the state of Chiapas. Specimens were collected in cloud forest, mixed oak-cloud forest, and mixed oak-pine forest, with some specimens collected on Alnus spp., Celtis spp., Chaetoptelea spp., Morella spp., Saurauia spp., and Quercus spp. Flight period was in January to August, November, and December (in both seasons). Previous altitudinal distribution records are between 88 and 2012 m [143,216,218] (based on geographical coordinates); herein, specimens were collected from 961 to 3088 m, which is the highest altitudinal distribution record for this species.
Remarks: Coniopteryx (C.) westwoodii presents a generally Nearctic distribution. This is the southernmost record for the species, ca. 961 km south from previous records (Ajutchitlán, Querétaro, Mexico). Specimens were collected in coffee plantations, cloud forest, and mixed oak-cloud forest. Flight period was in February to March, and December (mainly in the dry season). Previous altitudinal distribution records are between 189 and 559 m ( [218] based on geographical coordinates); herein, specimens were collected from 961 to 2454 m, which is the highest altitudinal distribution record for this species.
Remarks: Coniopteryx (S.) fumata was previously known from Central and South America. This is the northernmost record for the species, ca. 1080 km north from previous records (Turrialba, Cartago, Costa Rica). A single male specimen was collected in cloud forest. Flight time was in March (in dry season). Previous altitudinal distribution records are between 1000 and 1500 m [80]; herein, the specimen was collected at 1106 m.
Remarks: Coniopteryx (S.) isthmicola presents a Central American, Neotropical distribution, with a few Nearctic records. Specimens were collected in coffee plantations and cloud forest. Flight period was from February and March (mainly in the dry season). Previous altitudinal distribution records are between 670 and 865 m ( [16,144]; based on geographical coordinates); herein, specimens were collected between 958 and 966 m, which is the highest altitudinal record for this species.

Coniopteryx (Scotoconiopteryx) josephus Sarmiento-Cordero & Contreras-Ramos, 2019
Distribution: Mexico (Colima, Chiapas (new record), Morelos, Oaxaca) [8,11]. Remarks: Coniopteryx (S.) josephus is known only from Mexico. This is the southernmost record for the species, ca. 439 km south from previous records (Santa María Huatulco, Oaxaca, Mexico). A single male specimen was collected in evergreen tropical forest, with flight time in January (in the dry season). Previous altitudinal distribution records are between 88 and 940 m [8]; herein, the specimen was collected at 661 m.

Coniopteryx (Scotoconiopteryx) quadricornis Meinander in Meinander & Penny, 1982
Distribution: Brazil [11,32,145,215,217] and Mexico (new record) (Chiapas). Remarks: Coniopteryx (S.) quadricornis was previously known from Brazil. This is the northernmost record for the species, ca. 5389 km north from previous records (Rondônia, Brazil). Specimens were collected in evergreen tropical forest, coffee plantations, and cloud forest. Flight period was in February, March, and May (mainly in the dry season). Previous altitudinal distribution records are unknown; herein, specimens were collected from 684 to 1230 m.
Remarks: Conwentzia barretti presents a Nearctic and Neotropical distribution, with a wide distribution in the Nearctic region. Specimens were collected in cloud forest, oak forest, and pine forest, with some specimens collected on Quercus sp. and Alnus sp. Flight period was from January to August, and November to December (mainly in the dry season). Previous altitudinal distribution records are between 441 and 2896 m [9,16,32,143,206]; herein, specimens were collected between 1705 and 3277 m, the highest altitudinal distribution record for this species.
Remarks: Semidalis boliviensis presents a Neotropical distribution. Specimens were collected in evergreen tropical forest and coffee plantations. Flight period was in April, June, August, October, and December (mainly in the rainy season). A previous altitudinal distribution record is from 1100 m [143]; herein, specimens were collected between 680 and 749 m, which is the lowest altitudinal record for the species.
Remarks: Semidalis hidalgoana presents a Nearctic and Neotropical distribution. Specimens were collected in evergreen tropical forest, coffee plantations, and cloud forest, with some specimens collected on Inga sp. Flight period was in February to March, May to September, and December (mainly in the rainy season). Previous altitudinal distribution records are unknown; herein, specimens were collected between 677 and 1612 m.
Remarks: Semidalis manausensis presents a Neotropical distribution. Specimens were collected in a mixed oak-cloud forest. Flight period was from July to October, and December (mainly in the rainy season). Previous altitudinal distribution records are between 1500 and 2100 m [144]; herein, specimens were collected from 2076 to 2444 m, which is the highest altitudinal record for this species.

Semidalis problematica Monserrat, 1984
Distribution: Mexico (Chiapas (new record), Veracruz) [11,32,34]. Remarks: Semidalis problematica is known only from Mexico. Specimens were collected in evergreen tropical forest, cloud forest, and mixed oak-cloud forest, with some specimens collected on Lauraceae spp., Miconia spp., and Myriocarpa spp. Flight period was in January to December (in both seasons). Previous altitudinal distribution records of this species are unknown; herein, specimens were collected between 667 and 2436 m.

Semidalis soleri Monserrat, 1984
Distribution: Costa Rica, Mexico (Chiapas (new record), Veracruz) [4,11,34,144]. Remarks: Semidalis soleri is known from Mexico and Costa Rica. Specimens were collected in evergreen tropical forest and coffee plantations, with some specimens collected on Saurauia spp. and Miconia spp. Flight period was in January to February, and April to December (in both seasons). Previous altitudinal distribution records are between 1500 and 2000 m [80,144]; herein, specimens were collected from 673 to 799 m, which are considered the lowest altitudinal records for this species. Distribution: Costa Rica, Panama [11,150], and Mexico (new record) (Chiapas). Remarks: Biramus aggregatus was previously known from Costa Rica and Venezuela. This is the northernmost record for the species, ca. 956 km north from previous records (Estación Biológica Monteverde, Puntarenas, Costa Rica). Specimens were collected in cloud forest. Flight period was from January to June (mainly in the dry season). Previous altitudinal distribution records are between 1300 and 1540 m [150]; herein, specimens were collected between 1657 and 1712 m, which is the highest altitudinal distribution record for this species.

Genus Hemerobiella Kimmins, 1940
Hemerobiella sinuata Kimmins, 1940 Distribution: Ecuador and Mexico (Chiapas) [4,11,83]. Remarks: Hemerobiella sinuata is known from Ecuador and Mexico. A single male specimen was collected in cloud forest. Flight time was in October (in the rainy season). A previous altitudinal distribution record is from 1000 m [222]; herein, the specimen was collected at 1586 m, which is the highest altitudinal distribution record for this species.
Remarks: Hemerobius alpestris presents a Nearctic and Neotropical distribution. Specimens were collected in oak forest and pine forest, with some specimens found on Pinus sp. Flight period was in January, March to June, and August to December (in both seasons). Previous altitudinal distribution records are between 1219 and 3200 m [9,18]; herein, specimens were recorded from 3030 to 3789 m, which is the highest altitudinal distribution record for the species. Variation: Specimens of this species typically have body and wings with a reddish tinge, but some studied specimens had a much paler wing pigmentation.
Remarks: Hemerobius bolivari presents a Neotropical distribution, although it was introduced to Portugal (Palearctic region). Specimens were collected in cloud forest, oak forest, and pine forest. Flight period was in January, March to April, June to August, and October to December (in both seasons). Previous altitudinal distribution records are between 300 and 2800 m [18,228]; herein, this species was recorded from 1123 to 3166 m, which is the highest altitudinal distribution record for this species.
Remarks: Hemerobius discretus presents a Nearctic and Neotropical distribution. Specimens were collected in cloud forest, oak forest, and pine forest, with some specimens found on Alnus spp., Licaria spp., Pinus spp., and Roldana spp. Flight period was from January to December (in both seasons). Previous altitudinal distribution records are between 914 and 3000 m [9,18,232]; herein, specimens were collected between 1732 and 3580 m, which is the highest altitudinal distribution record for this species.
Remarks: Hemerobius domingensis presents a Neotropical distribution. Specimens were collected in cloud forest and oak forest, with some specimens found on Lauraceae sp. Flight period was in January, March to July, and September (mainly in the rainy season). Previous altitudinal distribution records are between 914 and 2133 m [18]; herein, specimens were collected from 1194 to 2438 m, which is the highest altitudinal distribution record for this species.
Remarks: Hemerobius gaitoi presents a Neotropical distribution. Specimens were collected in cloud forest and oak forest. Flight period was in January to September, and November to December (in both seasons). Previous altitudinal distribution records are between 870 and 2100 m [18]; herein, specimens were collected from 1155 to 2377 m, which is the highest altitudinal distribution record for this species.
Remarks: Hemerobius hernandezi presents a Neotropical distribution. Specimens were collected in evergreen tropical forest, cloud forest, and oak forest, with some specimens found on Justicia sp., Miconia spp., and Quercus spp. Flight period was from January to December (in both seasons). Previous altitudinal distribution records are between 90 and 2200 m [18]; herein, specimens were collected from 661 to 2205 m.
Remarks: Hemerobius hirsuticornis presents a Neotropical distribution. Specimens were collected in evergreen tropical forest, coffee plantations, and cloud forest. Flight period was in January and March (mainly in the dry season). Previous altitudinal distribution records are between 550 and 1500 m [80,222]; herein, specimens were collected from 743 to 1194 m.
Remarks: H. jucundus is known from Mexico and Central America. Specimens were collected in evergreen tropical forest, cloud forest, oak forest, and pine forest, with some specimens found on Alnus spp., Clethra spp., Fuchsia spp., Licaria spp., Pinus spp., and Roldana spp. Flight period was from January to December (in both seasons). Previous altitudinal distribution records are between 1219 and 2896 m [9,18]; herein, specimens were collected from 736 to 3358 m, which are the lowest and highest altitudinal distribution records for this species.
Remarks: Hemerobius martinezae is known from Mexico and Central America. Specimens were collected in cloud forest, oak forest, and pine forest, with some specimens found on Alnus spp. and Roldana spp. Flight period was in January to July, and September to December (in both seasons). Previous altitudinal distribution records are between 1219 and 2900 m [9,18]; herein, specimens were collected from 1470 to 3128 m, which is the highest altitudinal distribution record for this species.

Hemerobius nigridorsus Monserrat, 1996
Distribution: Costa Rica, Venezuela [11,18,228], and Mexico (new record) (Chiapas). Remarks: Hemerobius nigridorsus was previously known from Costa Rica and Venezuela. This is the northernmost record for this species, ca. 1220 km north from previous records (La Amistad International Park, Puntarenas, Costa Rica). Two male specimens were collected in cloud forest. Flight period was in April to May, and October to November (mainly in the rainy season). Previous altitudinal distribution records are between 1500 and 1600 m [18,80]; herein, specimens were collected from 1705 to 1712 m, which is the highest altitudinal distribution record for this species.
Remarks: Hemerobius withycombei presents a Neotropical distribution. Specimens were collected in evergreen tropical forest and coffee plantations. Flight period was in March, July, and August (mainly in the rainy season  [11,35,82,83,95,136,169,203,226,[235][236][237][238][239]. Remarks: Megalomus minor presents a Nearctic and Neotropical distribution. Specimens were collected in evergreen tropical forest, coffee plantations, and cloud forest, with some specimens found on Clibadium spp., Eupatorium spp., Inga spp., and Saurauia spp. Flight period was from January to September (mainly in the rainy season). Previous altitudinal distribution records are between sea level and 1500 m [95]; herein, specimens were collected from 657 to 1209 m.
Remarks: Megalomus pictus is known from Mexico and Central America; it was previously reported for Mexico, however a specific locality was unknown. Specimens were collected in mixed oak-cloud forest and pine forest. Flight period was in May, June, and September (mainly in the rainy season). Previous altitudinal distribution records are between 2200 and 3200 m [80,95,239]; herein, specimens were collected from 2081 to 3187 m, which includes the lowest altitudinal distribution record for this species.

Subfamily
Remarks: Micromus subanticus presents a Nearctic and Neotropical distribution. A single female specimen was collected in cloud forest. Flight time was in April (in the dry season). Previous altitudinal distribution records of this species are unknown; herein, a specimen was collected at 1479 m.
Remarks: Nusalala championi is known from Mexico and Central America. Specimens were collected in evergreen tropical forest, coffee plantations, cloud forest, and mixed oak-cloud forest, with some specimens found on Psidium spp. Flight period was in January to August, and October to December (in both seasons). Previous altitudinal distribution records are between 610 and 1524 m [80,96,246]; herein, specimens were collected from 775 to 2174 m, which is the highest altitudinal distribution record for this species. Variation: Individuals of this species typically have forewings with 5-7 radial sector branches, but some specimens had only four branches.
Remarks: Nusalala irrebita is known from Mexico and Central America. Specimens were collected in cloud forest and oak forest. Flight period was in March, May, July, October, and December (in both seasons). Previous altitudinal distribution records are between 1300 and 1600 m [80,96]; herein, specimens were collected from 1194 to 2452 m, which are the lowest and highest altitudinal distribution records for this species.
Remarks: Nusalala tessellata presents a Nearctic and Neotropical distribution. One female specimen was collected in evergreen tropical forest. Flight time was in June (in the rainy season). Previous altitudinal distribution records are between 250 to 2743 m [96,246]; herein, the specimen was collected at 722 m.

Nusalala unguicaudata Monserrat, 2000
Distribution: Mexico (Chiapas (new record), Nayarit), Costa Rica, and Guatemala [11,96]. Remarks: Nusalala unguicaudata is known from Mexico and Central America. Specimens were collected in evergreen tropical forest and coffee plantations, with some specimens found on Eupatorium spp. Flight period was in January to February, April, and December (mainly in the dry season). Previous altitudinal distribution records are from 1500 m [80,96]; herein, specimens were collected from 678 to 774 m, which are the lowest altitudinal records for this species.  [11,35,37,213,248], and Mexico (new record) (Chiapas).

Subfamily
Remarks: Notiobiella cixiiformis presents a Neotropical distribution. This is the northernmost record for this species. A single female specimen was collected in mixed oak-cloud forest. Flight time was in May (in the rainy season). A previous altitudinal distribution record is from 1000 m [35]; herein, the specimen was collected at 2060 m, which is the highest altitudinal distribution record for this species.

Notiobiella mexicana Banks, 1913
Distribution: Costa Rica and Mexico (Chiapas (new record), Jalisco, San Luis Potosí) [4,11,80]. Remarks: Notiobiella mexicana is known from Mexico and Costa Rica. Specimens were collected in evergreen tropical forest, with some specimens collected on Inga spp. Flight period was in April and May (in both seasons). Previous altitudinal distribution records are between sea level and more than 1000 m [80]; herein, specimens were collected from 670 to 693 m.

Subfamily Sympherobiinae Comstock, 1918 Genus Sympherobius Banks, 1905 Sympherobius axillaris Navás, 1928
Distribution: Mexico (Chiapas (new record), Ciudad de México) [11,31]. Remarks: Sympherobius axillaris is only known from Mexico (Nearctic). Herein, we record this species after 93 years of its original description. This is the southernmost record of the species, ca. 800 km south from previous records (Peñón Viejo, Mexico). Specimens were collected in cloud forest, oak forest, and pine forest. Flight period was from March to May, August to September, and November (in both seasons). Previous altitudinal distribution records for this species are unknown; herein, specimens were collected from 2406 to 3205 m.
Remarks: Sympherobius distinctus presents a Nearctic and Neotropical distribution. A single male specimen was collected in mixed oak-cloud forest. Flight time was in May (in the rainy season). A previous altitudinal distribution record is from 2750 m [97]; herein, the specimen was collected at 2060 m, which is the lowest record for this species.

Sympherobius marginatus (Kimmins, 1928)
Distribution: Guatemala [11,36], Mexico (new record) (Chiapas). Remarks: Sympherobius marginatus was previously known only from Guatemala. Herein, we record this species after its original description 92 years ago. This is the northernmost record for this species, ca. 82 km north from previous records (Cerro (Volcán) Zunil, Guatemala). Specimens were collected in cloud forest, oak forest, and pine forest. Flight period was from February to June (mainly in the dry season). Previous altitudinal distribution records are between 1220 and 1524 m [36]; herein, specimens were recorded from 1568 to 3176 m, which is the highest altitudinal record for this species. Variation: Individuals of this species typically have forewings with membranes that are dark brown, but some specimens had forewings with pale pigmentation.
Remarks: Sympherobius similis presents a Nearctic and Neotropical distribution. Specimens were collected in cloud forest and oak forest. Flight period was in March and April (in the dry season). Previous altitudinal distribution records are between 1000 and 1768 m [243]; herein, specimens were collected from 1168 to 2079 m, which is the highest altitudinal record for this species.

Sympherobius subcostalis Monserrat, 1990
Distribution: Mexico (Chiapas, Jalisco, Veracruz, Yucatán) [4,11,35,80]. Remarks: Sympherobius subcostalis is known only from Mexico (Neotropical). Specimens were collected in evergreen tropical forest and coffee plantations. Flight period was in January and September (in both seasons). Previous altitudinal distribution records are between sea level and 630 m [35,80]; herein, specimens were collected from 700 to 748 m, which is the highest altitudinal record for this species.

Sympherobius sp.
Distribution: Mexico (Chiapas). Remarks: A single male specimens was collected in pine forest, with flight time in May (in the rainy season), at an altitude of 3181 m. The specimen is morphologically similar to Sympherobius angustus Banks, 1904, and Sympherobius killingtoni Carpenter, 1940.
Remarks: Nolima infensa presents a Neotropical distribution. Specimens were collected in cloud forest, with some specimens collected on Clibadium spp. Flight period was in February and October (in both seasons). Previous altitudinal distribution records are between 396 and 1500 m [99]; herein, specimens were collected from 1250 to 1479 m.
Remarks: Nolima victor is known from Mexico and Guatemala (Nearctic and Neotropical). Specimens were collected in cloud forest. Flight period was in October and November (in both seasons). Previous altitudinal distribution records are between 244 and 2775 m [38,99]; herein, specimens were collected at 1479 m. Remarks: Dicromantispa sayi presents a Nearctic and Neotropical distribution. Specimens were collected in evergreen tropical forest, coffee plantations, and cloud forest, with some specimens found in Coffea spp. Flight period was in January and August to September (mainly in the rainy season). Previous altitudinal distribution records are between 11 and 1239 m [80,85,169,261]; herein, specimens were collected from 659 to 775 m. Variation: Individuals of this species typically have a a yellow-brown body coloration, but some specimens had a red-brown body coloration.
Remarks: Leptomantispa pulchella presents a Nearctic and Neotropical distribution. A single male specimen was collected in evergreen tropical forest. Flight time was in February (in the dry season). A previous altitudinal distribution record is from 1500 m [85]; herein, the specimen was collected at 694 m, which is the lowest altitudinal record for this species. Distribution: Mexico (Chiapas, Hidalgo, Jalisco, Quintana Roo, Sinaloa, Veracruz, Yucatán), Costa Rica, Guatemala, and Panama [4,11,39,80,83].

Genus
Remarks: Haploglenius flavicornis is known from Mexico and Central America (Neotropical). A single female specimen was collected in cloud forest, with a flight time in September (in the rainy season). Previous altitudinal distribution records are between 10 and 1520 m [80]; herein, the specimen was collected at 926 m.
Remarks: Ululodes bicolor presents a Nearctic and Neotropical distribution. A single male specimen was collected in evergreen tropical forest. Flight time was in July (in the rainy season). Previous altitudinal distribution records are between sea level and 500 m [80]; herein, the specimen was collected at 696 m, which is the highest altitudinal record for this species.

Ululodes sp.
Distribution: Mexico (Chiapas). Remarks: A single female specimen was collected in evergreen tropical forest, with flight time in November (in the dry season), at an altitude of 661 m. The specimen is morphologically similar to Ululodes cajennensis (Fabricius, 1787), but with evident differences in size, color patterns, and genitalia.
Remarks: Myrmeleon (M.) immaculatus presents a Nearctic and Neotropical distribution. This is the southernmost record for this species. Larval specimens, which were reared to adults, were collected in cloud forest and oak forest in February, March, May, December (mainly in the dry season), and one adult specimen in August (in the rainy season). Larval specimens became pupae between February and June, and emerged as adults between April and June. The altitudinal distribution of this species is unknown; in the present study, the altitudinal distribution was recorded from 955 to 1749 m.
Remarks: Myrmeleon (M.) timidus presents a Nearctic and Neotropical distribution. Larval specimens, which were reared to adults, were collected in evergreen tropical forest and coffee plantations in February and July (in both seasons). Larval specimens became pupae during February to April, August and November, and emerged as adults between February to May, August, September and November. Previous altitudinal distribution records are between sea level and at least 400 m [80,168]; herein, specimens were collected from 704 to 746 m, which is the highest altitudinal record for this species.
Remarks: Myrmeleon (M.) uniformis is known from Mexico and Central America (Nearctic and Neotropical). Specimens, some reared from larval stage to adult, were collected in cloud forest and oak forest in February (in the dry season), and other were collected in the adult stage from May to July (in the rainy season). Larval specimens became pupae during March, and emerged as adults in April. Previous altitudinal distribution records are from sea level to 1700 m [41,80]; herein, specimens were collected from 1514 to 2173 m, which is the highest altitudinal record for this species.

Keys to Families and Genera of Neuroptera from Volcán Tacaná, Chiapas, Mexico
See Appendix B for the Spanish version of the keys.

Key 2: Genera of Coniopterygidae (adult males and females)
(after [8,146] 5b. Genitalia with gonapsis absent (rarely with gonapsis); gonarcus and arcesus with no horn-like structures 7 6a. Abdomen with sternite 8 + 9 short and not fused; genitalia with gonarcal bridge wide, and gonapsis long (in relation to S8 + 9) with variable shape Ceraeochrysa 6b. Abdomen with sternite 8 + 9 elongate and fused; genitalia with gonarcal bridge narrow, and gonapsis short (in relation to S8 + 9), spoon-shaped Titanochrysa 7a. Head with scapes elongated or modified and/or with horns or cavities on the frons; genitalia with pseudopenis present Meleoma 7b. Head with scapes not elongated, and with no modifications, horns or cavities on the frons; genitalia with pseudopenis absent Chrysopodes

Altitudinal Distribution of the Neuropteran Fauna from Volcán Tacaná, Chiapas, Mexico
The fauna of Neuroptera has a wide distribution along the sampled altitudinal gradient (650-3360 m), and this general fauna was not divided in distinct groups across the altitudinal gradient; nevertheless, there is a tendency for the lower altitudes to share the same species (Figure 4a), with the largest number of lacewing species occurring at low and medium altitudes in the volcano. It is evident that two lower levels have a similar fauna of lacewings, sharing the presence of Ceraeochrysa achillea, C. sanchezi, C. squama, Leucochrysa

Altitudinal Distribution of the Neuropteran fauna from Volcán Tacaná, Chiapas, Mexico
The fauna of Neuroptera has a wide distribution along the sampled altitudinal gradient (650-3360 m), and this general fauna was not divided in distinct groups across the altitudinal gradient; nevertheless, there is a tendency for the lower altitudes to share the same species (Figure 4a Table S1); 0 = absence; 1 = presence.  Table S1); 0 = absence; 1 = presence.
Several species presented a wide distribution in the altitudinal gradient, two of them are present in all sampling altitudes-Hemerobius jucundus and Neoconis dentata-while Ceraeochrysa arioles, C. berlandi, Chrysopodes (C.) varicosus, Plesiochrysa brasiliensis, Semidalis problematica, Hemerobius hernandezi, and Nusalala championi are present in the altitudes between 600 and 2000 m. In addition, Coniopteryx (C.) simplicior and Hemerobius bolivari are distributed between 1000 to 3000 m, with no occurrence at lower altitudes.
A large number of species had a restricted distribution (Figure 4a). At the lowest altitude (ca. At the family level, different results were obtained on the relationship of the different altitudinal ranges. Chrysopidae, the family with the largest number of species in the Tacaná volcano, presents a similar distribution pattern to that observed for Neuroptera in general ( Figure 4b); nevertheless, for the whole order, the two lowest levels (ca. 600 and 1000 m, respectively) are closer to each other in species composition than either is to the medium level (ca. 1400 m), while the composition of chrysopid species is similar between the three lower levels. Eight genera of green lacewings were recorded from the volcano, of which Ceraeochrysa, Chrysoperla, Chrysopodes, Leucochrysa, and Plesiochrysa occurred in a wide range between 600 m to 2000 m, with Ceraeochrysa reaching higher altitudes. The three remaining genera were distributed between the middle and high altitudes, with Titanochrysa exclusively in the middle altitudes (between 1200 to 1600 m), Meleoma present between the middle and high altitudes (1200 to 3000 m), and finally Ungla restricted to high altitudes (between 2000 to 3000 m).
Hemerobiidae presented an inverse pattern to that observed for Neuroptera in general, i.e., most species were shared between the highest altitudinal ranges (Figure 4c). Eight genera of brown lacewings were reported for the Tacaná volcano, of which Hemerobius, Megalomus, Notiobiella, Nusalala, and Sympherobius have a wide range of altitudinal distribution (between 600 to 3000 m), with Hemerobius, Megalomus, and Sympherobius present at the highest altitudinal levels, contrary to other genera-Biramus, Hemerobiella, and Micromus-restricted exclusively to medium altitudes (1400 to 1700 m).
Myrmeleontidae have no species recorded at the highest level of the volcano (ca. 3000 m); however, when the other four sampled levels are compared, there is a tendency for the highest levels to share the same species (Figure 4d). Such family presented the lowest number of genera of all families, with a total of three genera, of which Ululodes and Haploglenius were restricted to low altitudes (600 to 900 m). On the contrary, Myrmeleon presented a wider altitudinal distribution, present from low to high altitudes (700 to 2000 m).
Mantispidae also had no species recorded at the highest level of the volcano (ca. 3000 m), and the species composition fluctuated along the altitudinal gradient, with most species occurring at the median altitudes (Figure 4e). Four genera of mantidflies were recorded. Zeugomantispa occurred mainly at high and medium elevations, with the highest distribution at 2000 m. In contrast, some genera were recorded at low altitudes (Dicromantispa and Leptomantispa) or medium altitudes (Nolima).
Coniopterygidae was present in all sampled levels; however, its species composition also fluctuated along the altitudinal gradient, without a definite tendency, similar to Mantispidae (Figure 4f). Only four genera of dustywings were recorded, of which Coniopteryx, Neoconis, and Semidalis presented a wide range in altitude (between 600 to 3000 m), with Coniopteryx and Neoconis present at the highest altitudinal levels. The genus Conwentzia was present from medium to high altitudes (1700 to 3000 m).
Rhachiberothidae was represented only by one species, Trichoscelia santareni, which is endemic to Mexico and was restricted to low and medium altitudes [85].

Biogeographic Composition of the Neuropteran Fauna from Volcán Tacaná, Chiapas, Mexico
Despite a fragmented knowledge about neuropteran distribution, herein, we attempt a characterization of the fauna from Tacaná volcano following the criteria of Morrone [280,281]. The fauna of lacewings had only 9% of the species of Nearctic affinity, with 16% of the species having a wide distribution in the New World. The largest portion of the fauna, ca. 75%, had a Neotropical affinity, with only Rhachiberothidae, 87% of Chrysopidae, 64% of Coniopterygidae, 72% of Hemerobiidae, and 50% of Myrmeleontidae species occurring in the Neotropical region. The recorded species of Mantispidae possess a wide distribution across the New World.
The Tacaná volcano is part of the Central American mountain range and its fauna has great affinity to the Neotropical region. More than 80% of the Tacaná volcano lacewing species occur in the Brazilian subregion, especially the Mesoamerican and Pacific domains, with approximately 50% and 71% of the species, respectively. Several neuropteran species (61%) are present only in the Mexican Transition Zone. Some species are also present in the Antilles and some subregions of South America. Most of the species with a Neotropical affinity are reported from the provinces of Puntarenas-Chiriquí and Guatuso-Talamanca (both Pacific)-which include around 50% of the species present in the Volcano-as well as from the Veracruzan, Pacific Lowlands, and Balsas Basin (all Mesoamerican), and the Transmexican Volcanic Belt and Chiapas Highlands (both from the Mexican transition zone). This last province received 52 new records of Neuroptera, of which 21 belong to the family Chrysopidae.
The Neuroptera fauna of the different altitudinal levels possess different biogeographic affinities, with lower elevational levels (650 m to 810 m) having a greater Neotropical affinity (79% of the species) and a smaller number of species of Nearctic affinity (5%). Nearctic affinity therefore increases with altitude, i.e., at the highest altitudes (2870-3360 m), 50% of the species have Neotropical affinity and 45% of them have Nearctic affinity.

Discussion
The number of studies about Neuroptera diversity is still low when compared with that about other insect orders; such studies are mainly focused on faunal lists of a particular country or region. Despite the Neotropical region being fourth in number of Neuropterida species [3], these Neuroptera species are still poorly known, which demonstrates a need for more studies in this area. The Nearctic region is sixth in the number of lacewing species [3]; however, this is the region with the largest number of studies about Neuroptera diversity in the Americas.
Several studies have recorded that neuropterans have a wide altitudinal distribution, from sea level to mountainous areas of more than 4000 m [2,80,282,283], which is supported in the present study, with neuropteran species recorded from 650 to more than 3500 m. Herein, we observed that neuropterans presented a higher species richness between 600 and 1700 m, with a few number of species occurring at altitudes above 3000 m, agreeing with the general information on the vertical distribution of the order; i.e., lacewings, although a cosmopolitan group, will individually present an affinity to tropical, subtropical, xeric and warm environments, or high mountain areas [3,284,285].
Some species, especially those from Chrysopidae, Hemerobiidae, and Coniopterygidae, were restricted to a single altitudinal range. It is noteworthy that the level with the highest number of species with restricted distribution was the lowest one (ca. 600 m), with a total of 19 species, while the highest level (ca. 3000 m) had the lowest number (4 spp.). Most of these restricted species represented new records for the state, the country, or potentially new species for science. This may be indicative of a possible high degree of endemism in the study site, and possible adaptations, in addition to the influence of biogeographical history, so it would be important to conduct studies focused on the ecological and biogeographical aspects that would explain these trends in the distribution of Neuroptera.
The Neuroptera fauna was biogeographically characterized, especially regarding possible biogeographical affinity (cf., [286]). The largest similarity of the neuropteran fauna at the Tacaná volcano occurred between levels 1 and 2 (between 600 and 1200 m); these levels were characterized by considerable anthropogenic impact as there were coffee plantations mixed with native vegetation (evergreen tropical forest or cloud forest). These areas also presented more Neotropical and cosmopolitan species, and as altitude increased, this composition was lost with the inclusion of taxa of Nearctic affinity. Overall, there is a strong affinity to the Pacific and Mesoamerican domains, supporting an evident Central and South American relationship of the fauna. This agrees with Halffter et al. [287], in that lineages of modern distribution constitute the typical Neotropical pattern, integrated after the consolidation of the Panamanian bridge with species close to those of northern South America, now distributed in the tropical lowlands of Mexico, or some representing the Mesoamerican mountain pattern, composed by taxa that evolved in the Central American Nucleus, often presenting expansions towards the north, and whose most important affinity is ancient South America.
Volcán Tacaná may be assumed to have a high potential for endemism, being part of the Central American Core Mountain system (within the Mexican Transition Zone subregion and the Chiapas Highlands province). Its neuropteran fauna is predominantly Neotropical with some species of Nearctic affinity and others of a wide distribution in the Americas; this conforms with Halffter et al. [288], in that mountains of the Mexican transition zone present a fauna with strong Nearctic affinity at their high altitudes (above 2400 m), while the fauna is related to the Neotropical region in the lowlands and plains (below 1200 m). The Nearctic affinity of the studied fauna is low when compared to other studies within the Mexican Transition Zone, such as Marquez-López et al. [9], who recorded the diversity of Hemerobiidae and Coniopterygidae in the Trans-Mexican Volcanic Belt, at ca. 2800 m, and obtained a higher Nearctic affinity. Based on this, the biogeographic affinities of Neuroptera may change notoriously, even between sites of the same subregion. Probably, the Neuroptera fauna is different in southern portions of the Mexican Transition Zone, due to a decrease in the number of species and lineages of northern origin, and the effect of the Isthmus of Tehuantepec that functions as a barrier, as was mentioned by Halffter [289], for entomofauna in general.
Altitudinal distribution patterns were different at the family level, which may be due to environmental conditions or habits that may sort the species out on the different levels. For instance, Chrysopidae and Hemerobiidae, despite presenting similar life histories [290], the former had a more diverse and shared species composition at low altitudes, while the latter displayed a similar pattern at medium and high altitudes.
The higher chrysopid species richness seen at low and medium altitudes was drastically lower at elevations above 3000 m; such diversity may be related to the presence of coffee plantations between 650 and 1770 m. This family is often present in agroecosystems, where they feed on various soft-bodied prey that are considered pests of different crops [137,291]. Most of the Chrysopidae species of the Tacaná volcano have a Neotropical affinity, which agrees with the literature [2,[292][293][294][295] about their large abundance and the diversification of green lacewings in tropical, subtropical, and temperate zones, and its lesser frequency in high and cold zones, with some species adapted to extreme temperature conditions.
Among the eight green lacewing genera recorded at the Tacaná volcano, five have a general distribution between 600 and 2000 m, with Ceraeochrysa and Leucochrysa showing a wider distribution compared to the other three (Chrysoperla, Chrysopodes, and Plesiochrysa). This broad altitudinal distribution has been previously recorded for these genera, with species recorded from sea level to more than 2000 m [12,13,80,106,137,138,140,296]. Both Ceraeochrysa and Leucochrysa, which presented the highest altitudinal range, displayed a high species diversity when compared to other genera, with two or three species reported to the region. Other genera were restricted to medium and high altitudes (Meleoma, Titanochrysa, and Ungla), which could be explained because these genera are associated with better-preserved environments, as opposed to low altitude sites with greater anthropogenic disturbance, such as coffee plantations where the other five genera, especially Ceraeochrysa, are often present [297].
Hemerobiidae displayed a higher number of species, most of them of Nearctic affinity, at middle altitudes between 1200 and 1700 m, decreasing at lower (ca. 600 m) and high altitudes (above 3000 m). Brown lacewings have a considerable affinity for oak and pine forests, as known from studies where Neuroptera present a strong affinity to the vegetation cover where they live [298,299]. Similar to Chrysopidae, this family presents substantial diversification in tropical, subtropical, and temperate environments, avoiding extreme environments such as deserts or high mountains [282]. Although both families have similar habits and distribution patterns, it is likely that ecological factors are influencing their distribution, perhaps with specific colonization mechanisms of brown lacewings at certain altitudinal levels to avoid possible competition for resources with Chrysopidae.
Eight genera of Hemerobiidae were recorded, five of them along the altitudinal gradient, from lowland areas characterized by evergreen tropical forest and coffee crops to high altitudes predominantly with pine forest. Species of Hemerobius, Megalomus, and Sympherobius have been previously reported with a broad altitudinal distribution, from sea level to over 3000 m [18,95,97,147,300,301]. Species of Nusalala and Notiobiella, known to have a wide altitudinal range, were not recorded above 2500 m, which could be due to the ecological requirements of these species, which have not been reported at altitudes above 1900 m [80,96,149], probably incapable of colonizing the highest peaks of the volcano.
Myrmeleontidae and Mantispidae had the smallest number of genera and species; despite previous records from a wide altitudinal range (sea level to ca. 3000 m) [80,85,99,265,266,302,303]; herein, both families had most species at low and medium altitudes (between 600 and 1500 m), with a decrease in the number of species up to 2000 m, and absence at altitudes higher than 3000 m. These families have been reported with higher species richness in tropical and warm environments, occurring less frequently or being absent in cold climates and high mountain areas [19,284,285], yet with few species reported between 2000 and 4000 m in Europe [304]. At the volcano, the maximum altitudinal record for these families was above 2000 m, with Myrmeleon (M.) uniformis and Zeugomantispa minuta. Genera such as Dicromantispa, Leptomantispa (Mantispidae), Ululodes, and Haploglenius (Myrmeleontidae) were only recorded at lower altitudes (from 600 to 900 m), while others such as Nolima (Mantispidae) remained at medium altitudes (1200 to 1500 m), which probably reflects that some genera are related to warmer environments with heterogeneous vegetation (as the low altitude sites studied here). Previous studies support that these families are typically present at low altitudes [244,298,[305][306][307].
Species of Coniopterygidae at the volcano were generally of Nearctic affinity. This family did not have a particular trend in its distribution, although in the highest areas above 3500 m its presence decreased considerably. This probably reflects that this family is mostly distributed in tropical and warm environments, and to a lesser extent in temperate zones or is absent in very cold climates [16,283,308]. Only four genera of dustywings were reported from the Tacaná volcano; three of them had a wide altitudinal distribution (Coniopteryx, Neoconis, and Semidalis), with Coniopteryx and Semidalis having previous records of a wide altitudinal distribution, while Neoconis has been reported at ca. 1600 m [16,43,143]. Conwentzia was present between 1700 and 3000 m, without occurring at low altitudes, and represented by a single species that has previous records between 400 and 2800 m [16].
Concerning the Rhachiberothidae, the only species collected in the volcano-Trichoscelia santareni-is endemic to Mexico, and has a Neotropical affinity, with records in the states of Tabasco and Quintana Roo [85].
This study highlights the importance of recording the distribution pattern of Neuroptera species for a better understanding and potential management of insect communities in protected areas, such as the Tacaná volcano. Noteworthy is the large number of new records of Neuroptera species for Mexico, which increases the known distribution range of such species and genera. Despite the Neuroptera fauna of the Tacaná volcano is most similar in lower altitudes, each studied family possessed a particular distribution and affinity along the altitudinal levels. The biogeographic affinity of the Neuroptera fauna from Tacaná volcano is variable between the altitudinal levels, with the lowest altitudes presenting species with Neotropical affinity, whereas higher levels have an increase in the number of species with Neartic affinity.      1a. Antenas apicalmente clavadas, usualmente tan largas como el cuerpo, pero algunas veces tan cortas como la cabeza y tórax combinados; alas anteriores y posteriores con una celda no alargada detrás del punto de fusión de Sc y RA (Ascalaphinae)   2 1b. Antenas apicalmente no clavadas, y no tan largas como la cabeza y tórax; alas ante-riores y posteriores con una celda alargada detrás del punto de fusión de Sc y RA 1a. Cabeza con vértex en forma de domo en vista frontal; patas anteriores con dos uñas pretarsales con ariola presente (Calomantispinae) Nolima 1b. Cabeza con vértex concavo en vista frontal; patas anteriores con una uña tarsal, con arolia ausente (Mantispinae)   2 2a. Pronoto, en vista lateral, con prominentes sedas en toda su longitud 3