A revision of Chilicola (Heteroediscelis), a subgenus of xeromelissine bees (Hymenoptera, Colletidae) endemic to Chile: taxonomy, phylogeny, and biogeography, with descriptions of eight new species

Abstract The bee subgenus Chilicola (Heteroediscelis) Toro & Moldenke, 1979 (Hymenoptera, Colletidae, Xeromelissinae) is revised. The subgenus is considered endemic to Chile and occurs across a broad range of habitats. Eight new species are described: Chilicola (Heteroediscelis) charizard Monckton, sp. n., Chilicola (Heteroediscelis) curvapeligrosa Monckton, sp. n., Chilicola (Heteroediscelis) guanicoe Monckton, sp. n., Chilicola (Heteroediscelis) katherinae Monckton, sp. n., Chilicola (Heteroediscelis) lickana Monckton, sp. n., Chilicola (Heteroediscelis) mayu Monckton, sp. n., Chilicola (Heteroediscelis) packeri Monckton, sp. n., and Chilicola (Heteroediscelis) randolphi Monckton, sp. n. One of the existing species, Chilicola (Heteroediscelis) valparaiso Toro & Moldenke, 1979, syn. n., is treated as a junior synonym of Chilicola (Heteroediscelis) mantagua Toro & Moldenke, 1979, and the nine remaining valid species are redescribed. Thoroughly illustrated keys to species for males and females are provided, along with habitus images, images of male terminalia, distribution maps for each species, and a map of relevant Chilean biogeographic regions. Results of phylogenetic analyses are presented, based upon 74 morphological characters and on CO1 barcode sequences, analyzed both separately and as a combined dataset. Monophyly of the subgenus is supported, and groupings within the subgenus are discussed in light of a biogeographic analysis of their species distributions (spatial analysis of vicariance), whereby divergence between taxa is found to occur primarily via north-south disjunctions.


Introduction
Chilicola Spinola, 1851 is a diverse neotropical genus of stem-nesting bees, comprising 98 named species (Ascher and Pickering 2015) in 15 subgenera (Packer 2008). The subgenus Chilicola (Heteroediscelis)  is notable in that its 10 named species are endemic to Chile (Packer 2008), where they occupy a broad range of habitats, from the hyper-arid Atacama Desert in the north, to moist temperate forests in the south, occurring at elevations from sea level to over 3200 m a.s.l.. Bees of the subgenus Heteroediscelis are small and slender, 4-7 mm long, and hylaeiform in appearance. They are mostly dark brown or black, with females having variable yellow markings on the legs and ventral surface of the antennae, and males having more extensive yellow colouration, particularly on the face. In males, the distinctly expanded hind legs and the robust, truncate process extending ventrally from the first sternum (S1 process) together readily distinguish this subgenus from all other Chilicola.
The subgenus Heteroediscelis was first described with 13 species . Three of these species were later moved to C. (Anoediscelis) Toro & Moldenke by Michener (1995) due to obvious differences: their males lack the combination of expanded hind legs and truncate S1 process characteristic of the other ten species. Toro and Moldenke (1979) recognized these differences but did not classify the three species in a different subgenus, citing the lack of corresponding differences in females as justification. Michener (1995) moved the other ten species to C. (Oediscelis) Philippi. Packer (2008) performed a phylogenetic analysis of Chilicola with exemplars from all known subgenera plus additional outgroup taxa, and found very strong support for the monophyly of Heteroediscelis, along with moderate support for its placement as sister to a well-supported clade formed by C. (Chilioediscelis) Chilicola s. str. Spinola, 1851; consequently, he resurrected the subgenus.
A number of new species belonging to Heteroediscelis have been recognized in collections for some time, several of them from geographic regions novel to the subgenus. However, no species-level taxonomic work has been done on the group since its description, and its phylogeny is unknown. Here, I synonymise one of the original species, redescribe the remaining nine and describe eight new species, bringing the total within the subgenus to seventeen. I also present a thorough phylogenetic analysis for the subgenus, as well as a discussion of the historical biogeography of these interesting little bees.

Taxonomy
For the revision, 1316 specimens of Heteroediscelis were examined. Of these, 344 were collected during field excursions over the course of this study, and the remaining 972 were obtained from five collections: the Packer Collection at York University, Toronto, Canada (PCYU), the American Museum of Natural History, New York, U.S.A. (AMNH), the collection at the Pontificia Universidad Católica de Valparaíso, Chile (PUCV), the Bee Biology and Systematics Laboratory, Logan, UT, U.S.A. (BBSL), and the collection at the Universidade Federal do Rio de Janeiro, Brazil (UFRJ). In addition to these collections, a number of specimens from PCYU will also be deposited at the Central Texas Melittological Institute (CTMI) upon completion of this work. As all specimens were collected in Chile, all locality information is written with country excluded.
Specimens were identified using the existing key  and those not matching the descriptions of the documented species were sorted into putative OTUs (operational taxonomic units) based on morphological characters. Sorted specimens were initially compared with paratype material of existing species where possible (later to holotypes for all species). Subsets of specimens from these sorted groups were selected for DNA barcoding (Hebert et al. 2003), particularly where either barcode sequences for that species were apparently not already recorded, or where specific identity was uncertain and comparison could be made to previously databased sequences.
For barcoding, a midleg from each specimen was removed and placed into a single well of a 96-well plate along with three drops of 95% EtOH. Extraction, PCR, and sequencing of the cytochrome c oxidase subunit I (CO1) were all done at the Canadian Centre for DNA Barcoding (CCDB) in Guelph, Canada, using standardized protocols (see Hebert et al. 2003, Ivanova et al. 2006. Successful output sequences were uploaded to the Barcode of Life Data Systems database (BOLD) (Ratnasingham and Hebert 2007). In order to check putative identifications against CO1 sequence data, barcode sequences were investigated in two ways: first, BOLD's built-in analysis platform was used to generate neighbour-joining trees from DNA barcodes based on the Kimura twoparameter (K80) base substitution model (Kimura 1980); second, all barcode sequences over 300bp were assigned Barcode Index Numbers (BINs) representing groups of similar sequences automatically generated by BOLD using a clustering algorithm (Ratnasingham and Hebert 2013). Sequences belonging to different BINs or to different groups on the neighbour-joining tree can subsequently be checked for morphological differences.
Accordingly, barcode sequences were used in species delimitation employing an integrative approach (e.g. Gibbs 2009a, 2009b, DeSalle et al. 2005, Meier et al. 2006. Identified BINs were investigated at the specimen level for any morphological differences that might support further separation into distinct groups, and additional specimens were selected for barcoding as necessary for a representative range of morphological variation and geographic coverage. Throughout, morphological characters were given precedence over DNA barcodes, as clear morphological differences were in some cases obvious among members of the same BIN (an outcome also noted by Gibbs 2009b). Groupings consistently supported by morphological characters were considered to be distinct species (i.e. using the phylogenetic species concept; Nixon and Wheeler 1990). Names assigned to groupings representing existing species were confirmed following examination of holotype material at AMNH.
Body measurements were taken using a calibrated ocular micrometer mounted on a Nikon SMZ1500 stereomicroscope. Measurements of body length, head width, thorax width, and forewing length were recorded in millimeters (mm). Due to distortions of the mesothorax from pinning these small bees, the standard measure of intertegular width could not be accurately assessed, and thorax width was instead measured across the pronotal lobes. Because the tagmata were not always in the same plane, body length was measured in sections (head, meso-and metasoma) and summed. To facilitate comparisons, measurements of pubescence are given relative to the diameter of the median ocellus (OD). Surface sculpture is described with puncture density stated in terms of puncture diameter (d) and interspace (i). All other measurements are expressed as ratios, with values representing ocular units at a common magnification of 90, where one ocular unit equals approximately 0.0137mm; these values have an approximate measurement error of +/-0.5 units. For measurements of the pedicel and antennal flagellomeres, greater precision was required, and these measurements were verified at 112.5 magnification (one ocular unit equals approximately 0.0110mm); their measurement error is estimated as +/-0.25 units.
Specimen records not originally having GPS coordinates are supplemented with them where possible, as determined via discussions with the collectors and using Google Earth (2013). Supplemented GPS coordinates are rounded to three decimal places and italicized in the descriptions, and include elevations recorded from the nearest road or path according to Google Earth. Specimens with barcode reference numbers or index/database numbers have these codes listed where applicable; multiple codes for a single specimen are listed separated by a double forward slash (i.e. //), and when multiple specimens are listed for a single locality, these codes are given in respective order, separated by commas. Each specimen was given a unique ID number (e.g. C.kate.001) in order to allow explicit reference to individual bees; these number codes are listed along with all specimen records in Suppl. material 2.
The following acronyms are used: OD: diameter of the median ocellus; MOC: distance from lower tangent of median ocellus to apex of clypeus; OOC: ocellocular distance, shortest distance between lateral ocellus and compound eye; IOC: interocellar distance, shortest distance between lateral ocelli; UOD & LOD: upper & lower interocular distance, shortest distance between inner orbits of compound eyes, above & below level of emargination, respectively; IOD: maximum interocular distance, or greatest distance between inner orbits of compound eyes at level of emargination; LOT: lower ocular tangent, an imaginary line crossing lower extremities of compound eyes in frontal view, perpendicular to main axis of face; IAD: interalveolar distance, shortest distance between antennal sockets; AOD: minimum antennocular distance, shortest distance between antennal socket and compound eye, typically at an angle of approximately 30° below horizontal laterad from antennal socket; F: antennal flagellomere; T: tergum; S: sternum.
Descriptions use terminology from Harris (1979) for surface sculpture and standard melittological terminology from Michener (2007) for structure, with the exception of the following non-standard terms: stria refers to a raised, rather than impressed, lineation; vertexal area refers to the dorsalmost surface of the head, between the compound eyes, frontal area, and occiput; clypeal lateral refers to the lateral, ventrally-bent portion of the clypeal apex; genal beard refers to the long erect hairs of the genal area; stigmal perpendicular refers to an imaginary line through the apex of the stigma and perpendicular to the costal margin of the forewing (Packer 2008); ventral metatibial convexity refers to the apicoventral expansion of the male metatibia, which has a read-ily defined summit; posterior metatibial concavity refers to the wide, deep longitudinal groove which extends the length of the male metatibia on the posterior surface; ventral/ posterior metatibial carina refers to the carinae associated with the posterior concavity of the male metatibia; apical metatibial lamina refers to the structure extending mediad from the base of the male metatibial spurs; metapostnotum refers to the dorsal surface of the propodeum (sensu Brothers, 1976). The following assumptions are made: when describing colouration, orange in place of normal yellow colouration is assumed to indicate discolouration due to exposure to cyanide, and is described as yellow for discoloured specimens; when describing legs, they are assumed to be fully extended and at right angles to the longitudinal axis of the body. Finally, frontal view is used to refer specifically to the angle of observation at which the lower tangent of the median ocellus and the apex of the clypeus are in a common plane.
All specimen images were taken using a Visionary Digital lift-operated imaging system and a Canon 5D Mark II digital SLR camera. Composite images were assembled using Helicon Focus stacking software. Images were edited, cropped, and scale bars added using Adobe Photoshop CS6 Extended ver. 13.0. Photographs marked in the caption with a " †" have been flipped horizontally to ease comparison. All images of new species are of the holotype (male) or allotype (female) except where otherwise indicated in the captions.
Specimens were examined for morphological characters that were potentially phylogenetically informative, and additional characters were adapted from the key and diagnoses of Toro and Moldenke (1979), and from the works of Gibbs and Packer (2006), Willis and Packer (2007), and Packer (2008). A total of 74 morphological characters were coded for each species, comprising 54 discrete and 20 continuous characters, the latter representing ratios between measured structures. Care was taken not to select inter-dependent characters; for example, characters were only included for both sexes when they did not vary in a consistent manner between males and females, thereby providing different phylogenetic information.
Leg colour characters in both sexes were considered to be potentially confounded, based on the observation that some combinations of colouration are never observed. In other words, certain changes in colour of different leg regions may occur together in single evolutionary steps. Non-redundant linear coding (NRLC; O'Grady and Deets 1987) of leg colour characters was therefore used to reduce fourteen potentially confounded characters to three pairs of multi-state characters representing the full range of leg colour variation (see Appendix 3 for explanation).
To ensure independence between continuous morphological characters, regression analyses were performed both on pairs of characters suspected to covary, as well as on male and female characters representing the same structure. Using this approach, adapted from Koch et al. (2015), characters were treated as dependent when they were found to have a significant linear regression without any outliers, indicating that they provide equivalent phylogenetic information -i.e. the variation in one character reliably predicts that of the other for all species. Conversely, the presence of an outlier for even one species is evidence that the two characters offer different phylogenetic information and are independent. Therefore, the magnitude of the difference between observed and expected values was calculated for each data point, and a threshold of 15% was used to identify outliers (arrived at through qualitative comparison of residuals between probable pairs of dependent and independent characters). For pairs of characters found to be dependent, the less variable character was discarded in order to capture maximal variation among species. For traits coded as discrete in one sex and continuous in the alternate sex, independence was assessed using continuous measurements taken from both sexes; if a significant linear regression was found with no outliers differing more than 10% from expected values, the continuous character in the alternate sex was discarded in favour of the discrete character. Finally, continuous characters were re-scaled to unity to avoid biases from magnitude (e.g. Koch et al. 2015).
Representative CO1 sequences were aligned in BOLD with default settings and using ClustalW. The sequence trace files were inspected in BOLD and sequences were edited if applicable -occasionally, BOLD truncates one or both ends of a sequence if it encounters a poor-quality read for a single base pair separating the tail from the rest of the sequence, even when the base pair reads beyond it are high-quality themselves. In such cases, excluded nucleotides were edited back into the full sequence for analysis. In all, this resulted in three datasets for phylogenetic analyses in TNT (Goloboff et al. 2008): morphological data only, molecular data only, and a combined dataset of both.
Maximum parsimony analysis was performed using extended implied weighting (Goloboff 1993, Golobof et al. 2008, Goloboff 2014 whereby discrete, continuous, and molecular data were treated as separate partitions in each analysis; the concavity value K was calculated independently for each partition using the "setk.run" script (as in Santos et al. 2014). The following values were used: discrete morphological, K = 5.78125 (N = 15.03); continuous morphological, K = 0.800781 (N = 15.05); molecular, K = 3.046875 (N = 14.86). Continuous characters are treated as additive by TNT (Goloboff et al. 2006) and characters derived from NRLC were also coded as additive (see Appendix 3). For all three analyses, a tree search was run for 100 replicates, using a driven search with default settings for ratchet, drift, and tree-fusing. The output was visualized using WinClada (Nixon 2002) and characters were mapped using unambiguous optimization, except where noted (see Suppl. material 1 for character-annotated tree, Fig. S1). Tree support is reported by fit, consistency index (CI), and retention index (RI). Support for nodes of the most parsimonious tree was assessed in TNT using absolute frequencies for bootstrap (BS) and GC values from symmetric resampling, both with 1000 replications (Goloboff et al. 2003). GC values represent the percentage of times that a given grouping was found at a node minus the number of times that the next most frequent arrangement was found without that grouping. 2 (1) Apical surface of S1 process longitudinally concave throughout ( Fig    Male: A C. mavida, malar space longer than clypeal lateral B C. mavida, S1 process † , lateral view with arrow indicating acute anterior apical angle C C. curvapeligrosa, malar space subequal to clypeal lateral D C. curvapeligrosa, S1 process † , lateral view with arrow indicating rounded apex; scale bars 0.25 mm.  9C); in lateral view, apex of S1 process appearing broadly rounded (Fig. 9D) ....... 10 10(8) First flagellomere subequal in length and width or shorter (<1.05×) than wide (Fig. 10A); clypeus extending for half its length or more below lower ocular tangent (Fig. 10A); malar space subequal to or slightly shorter than clypeal lateral (Fig. 10B); clypeal apex yellow along entire width (Fig. 10B)   Male: A C. curvapeligrosa, frontal view, F1 shorter than wide, clypeus extending for more than half its length below LOT (white portion of bracket) B C. curvapeligrosa, malar space subequal to clypeal lateral C C. mayu, frontal view, F1 ~1.1× as long as wide, clypeus extending for less than half its length below LOT (white portion of bracket) D C. mayu, malar space ~0.8× as long as clypeal lateral; scale bars 0.25 mm. 11(5) Malar space longer than (>1.05×) clypeal lateral (Fig. 11A); in lateral view anterior margin of S1 process strongly concave (Fig. 11B); metatibia with wide yellow basal ring (up to one fifth of tibial length) and apex yellow only in ventral half (Fig. 11C), sometimes very narrowly yellow in dorsal half ....... 12 -Malar space shorter than (<0.95×) clypeal lateral (Fig. 11D); in lateral view anterior margin of S1 process sublinear apically (Fig. 11E); metatibia more extensively pale, with wide (up to one third of tibial length) yellow basal and apical rings ( Male: A C. katherinae, malar space longer than clypeal lateral B C. charizard, S1 process, lateral view showing strongly concave anterior margin C C. charizard, metatibia with wide yellow basal ring and apex yellow only basally (C.chzd.005) D C. randolphi, malar space shorter than clypeal lateral E C. deserticola, S1 process † , lateral view showing sublinear anterior margin F C. deserticola, metatibia with basal third yellow and wide yellow apical ring; scale bars 0.25 mm.
Colouration: Black-brown except as follows: antenna variable (scape with yellow apicoventral spot to entirely dark; apicoventral rim of pedicel brown to entirely dark; ventral surface of flagellum yellow to yellow-brown, narrowing basally on F1, remaining flagellomeres suffused with brown basally). Following parts yellow: labrum; clypeus except narrowly to broadly dark along epistomal suture and apex usually brown laterad; lower paraocular area extending ~1OD above transverse portion of epistomal suture and not reaching lower tangent of antennal socket; protrochanter apical rim posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; protibia except dark medioventrally; probasitarsus, at least on dorsal half; prodistitarsus, at least on apical two-thirds; narrow apical ring on mesofemur; wide basal ring and narrow apical ring on mesotibia, the latter wider anteriorly; apical ring on metafemur, broad dorsally; broad basal and apical rings on metatibia, excluding apical margin of ventral convexity; metabasitarsus in basal quarter dorsally, extending to apex ventrally; anterior spot on tegula. Apicoventral rim of metacoxa yellow-brown. Ventral metatibial carina black. Metasoma dark brown, T1-T6 marginal zones translucent yellow.
Colouration: Black to black-brown except as follows: mandible with yellow basal spot, translucent yellow in apical half except apex translucent brown; antenna variable (apicoventral margin of F3 yellow-brown to brown; ventral surface of flagellum from F4 to terminal flagellomere yellow-brown to yellow, flagellomeres suffused with brown basally). Following parts yellow: dorsal surface of protibia, darkened to yellow-brown apicodorsally; apical half of prodistitarsus; narrow basal ring on mesotibia, wider dorsally. Anterior spot on tegula absent or faintly yellow. Following parts brown: apicodorsal spot on mesotibia; apical one-third of mesodistitarsus; narrow apical ring on metafemur (yellow-brown on some specimens). Metatibia with wide basidorsal band yellow-brown. Metasoma black, T1-T5 marginal zones narrowly translucent yellow.
Colouration: Black-brown, following parts yellow: labrum; clypeus except broadly dark along epistomal suture and apex brown laterad; lower paraocular area, extending further medially than laterally, more than 1OD above transverse portion of epistomal suture, not reaching lower tangent of antennal socket; apicoventral spot on scape; apicoventral surface of pedicel; ventral surface of F1-F5, F5 suffused with brown; apical rim of protrochanter posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including dorsal half of anterior surface; narrow apical ring on mesofemur; narrow basal ring and apicodorsal rim of mesotibia; basal quarter of metatibia, as well as apical quarter in ventral half only; basal quarter of metabasitarsus ventrally; anterior spot on tegula. Ventral surface of F6-F10 yellow-brown. Ventral surface of F11 brown. Apicoventral rim of metacoxa brown. Metasoma black, T1-T6 marginal zones amber to translucent yellow at margins.
Colouration: Black to black-brown except as follows: mandible with yellow basal spot, translucent yellow in apical half except apex translucent brown. Following parts brown: pedicel, F1 and F2, on apicoventral margins (the latter two additionally with small yellow-brown apicoventral spot); apicodorsal rim of metafemur; basidorsal spot on metatibia. Following parts yellow: ventral surface of antenna from F3 to terminal flagellomere; dorsal surface of protibia; apical half of prodistitarsus; basidorsal and apicodorsal spots on mesotibia; anterior spot on tegula. Metasoma black, T1-T5 marginal zones narrowly translucent yellow.
Variation. Some females have the apicoventral surface of pedicel, F1 and F2 yellow-brown, and/or a brown or yellow-brown spot on the lower paraocular below the anterior tentorial pit.
Etymology. The specific epithet is in homage to the fictional monster which this species resembles. It is treated as a noun in apposition. Diagnosis. Males are diagnosable by the combination of S1 process in lateral view having a rounded anterior apical angle, malar space subequal to or slightly shorter than the clyperal lateral, and first flagellomere no longer than wide. Females are diagnosable by the combination of malar space subequal in length to the clypeal lateral (0.9-1×)and leg colouration with apex of metatibia black-brown and following parts brown: apicodorsal spot on mesotibia, apicodorsal rim of metafemur, basidorsal spot on metatibia. Females of C. mavida are most similar, but have lighter leg colouration (apicodorsal spot on mesotibia yellow-brown, apicodorsal rim of metafemur yellow, wide basal ring of metatibia yellow, posterior apical rim of metatibia yellow-brown).
Colouration: Black-brown, following parts yellow: labrum; clypeus except broadly dark brown along epistomal suture and apex yellow along entire width; lower paraocular area, extending ~1OD above transverse portion of epistomal suture medially, further laterally but not reaching lower tangent of antennal socket; apicoventral surface of scape; protrochanter apical rim posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including dorsal half of anterior surface; narrow apical ring on mesofemur; basal and apical ring of mesotibia, the latter extending to cover apical half of anterodorsal surface; apicodorsal rim of metafemur; ventral surface of metatibia, as well as dorsal surface in basal quarter, narrowly yellow apically except broad around apical lamina and posterior concavity; basal half of metabasitarsus ventrally; anterior spot on tegula; apex of T7. Following parts orange-brown: ventral surface of antenna from apical half of pedicel to terminal flagellomere, narrowing basad on pedicel and F1; ventral surface of mesobasitarsus. Apicoventral rim of metacoxa and metatrochanters translucent yellow. Apical half of prodistitarsus brown. Metasoma black, T1-T6 amber marginal zones to translucent at margins.
Colouration: Black to black-brown except as follows: mandible with yellow basal spot, translucent yellow in apical half except apex translucent brown. Following parts brown: apicoventral margin of pedicel; apical two-thirds of prodistitarsus; apicodorsal spot on mesotibia; apicodorsal rim of metafemur; basidorsal spot on metatibia. Following parts yellow-brown: ventral surface of antenna from F3 to terminal flagellomere, flagellomeres suffused with darker brown basally; posterior surface of mesobasitarsus. Following parts yellow: dorsal surface of protibia; narrow basal ring on mesotibia; anterior spot on tegula. Metasoma brown, T1-T5 marginal zones amber to translucent yellow at margins.
Type material ( Variation. Some males have the clypeal apex brown laterad. Some females have the apicoventral margin of F1 and F2 brown or yellow-brown, and/or a yellow-brown or brown spot on the lower paraocular area below the anterior tentorial pit. Many females have an orange-brown metasoma with the gradulus and usually disc of T1-T6 brown, or at minimum darker than the pregradular and marginal zones. Males from Cerro El Roble and Espinalillo have the F1 longer than wide, as in C. mavida, but resemble C. curvapeligrosa in both colouration and malar space length; females from these localities are consistent with the diagnosis of C. curvapeligrosa. Distribution. Southern Andean from Farellones (RM Santiago) south to Paso Vergara (Region VII), also known from Cerro El Roble and Espinalillo (RM Santiago) in the Central Coastal Cordillera; 877-3026m a.s.l.
Etymology. The specific epithet is a nounphrase inspired by the signs encountered on the road leading to the type locality; it refers to the hairpin turns used to achieve high elevation. It is to be treated as a noun in apposition.
Comments. Specimens from Cerro El Roble and Espinalillo may belong to a divergent population, distinct from those found at higher altitudes in the Andes, and are thus excluded from the type series (see variation, above). Diagnosis. Males are diagnosable by the combination of malar space short (<0.5× clypeal lateral), mesobasitarsus mostly yellow, and metatibia with the combination of ventral carina extending full length of longitudinal concavity and toothed, posterior carina absent basally and originating in apical half of ventral carina, and apical lamina reduced (~0.5OD in length). Females are diagnosable by the combination of the malar space short (~0.4× clypeal lateral), frontal area without longitudinal depressions which partially accommodate the scape, and clypeus evenly and moderately densely punctate throughout (i=1-2d).
Colouration: Black-brown, following parts yellow: labrum; clypeus except narrowly dark along epistomal suture near anterior tentorial pit and apex usually brown laterad; lower paraocular area extending ~OD above transverse portion of epistomal suture medially, further laterally and nearly reaching lower tangent of antennal socket; apicoventral surface of scape; apical one-third of profemur on anterior surface, apical quarter on posterior surface; protibia except dark medioventrally; protarsus; broad apical ring on mesofemur, wider anteriorly; broad basal and apical rings on mesotibia, expanded to basal and apical one-third or continuous on dorsal margin; mesobasitarsus ventrally to entirely; wide apical ring on metafemur; basal quarter and apical quarter of metatibia, apical colouration extending to summit of ventral convexity; metabasitarsus ventrally and at least basal one-third dorsally; anterior spot on tegula. Following parts yellow-brown: apicoventral surface of pedicel; ventral surface of antenna from F1 to terminal flagellomere, narrowing basally on F1; protrochanter apical rim posteriorly; apical one-third of mesodistitarsus. Apicoventral rim of metacoxa and metatrochanter brown. Ventral metatibial carina black. Metasoma dark brown, T1-T7 marginal zones yellow-brown to translucent yellow at margins.
Colouration: Black to black-brown except as follows: mandible with yellow basal spot, otherwise translucent yellow to yellow except apex translucent brown. Following parts brown: apicoventral margin of pedicel and F1; posterior apical rim of metatibia (yellow-brown on some specimens). Following parts yellow: apicoventral surface of F2; ventral surface of antenna from F3 to terminal flagellomere, suffused with yellowbrown basally; dorsal surface of protibia, narrow basal and apical rings; probasitarsus; apical two-thirds of prodistitarsus; wide basal ring and narrow apical ring on mesotibia; mesobasitarsus ventrally; ventral half of metabasitarsus; apical half of metadistitarsus brown; anterior spot on tegula. Following parts yellow-brown: narrow apical ring on metafemur; wide basal ring on metatibia (yellow on some specimens). Metasoma black, T1-T5 marginal zones yellow-brown to translucent yellow at margins.
Structure: Labrum ~2.5× wider than long (19.5:7.5); malar space ~0.4× as long as clypeal lateral (3:7); LOT below anterior tentorial pits; clypeus subequal in length to maximum width in frontal view (24:26) extending for less than half of its length beyond LOT; median longitudinal groove weak on clypeus; subantennal sutures subequal in length to the shortest distance between them (11.5:11); IAD ~1.6× AOD (11.5:7); scape ~3.33× as long as maximum width (20:6); pedicel longer than wide (8:6.5); F1 subequal in length and width (5.5:5); F2 ~0.67× as long as F1 (3.5) and shorter than F3  Comments. The type series (male holotype & one male paratype) is labeled Atacama, Pueblo Hundido, x.1972, L. Ruz, on Heliotropium stenophyllum. Atacama is the name of Region III, thus the locality is interpreted as Diego de Almagro (formerly called Pueblo Hundido), and was indicated as such on a map of type localities by Toro and Moldenke (1979). However, besides the type series, C. deserticola has not been collected from further north than Carrizal Bajo, Region III, and H. stenophyllum does not occur north of Totoral, Region III (Luebert 2013), while Diego de Almagro is located 1.1° latitude further north than Totoral and in a different biogeographic region (Peña 1966, O'Brien 1971, Artigas 1975. In contrast, Pueblo Hundido is also the name of a village in Region IV (Coquimbo) to the southeast of Vicuña (COPEC 2012, Sernatur 2012); a road sign marks its location at approximately S 30.0629° W 70.4911° (Google Earth 2013). This village is within the range of H. stenophyllum (Luebert 2013) and is the apparent provenance of a specimen collected on that host plant species, labeled Coquimbo, Pueblo Hundido, x.1972, V. Cabezas. Specimen records show that Ruz, Toro, Neff, and Cabezas traveled together in September/October 1972 (see Supplementary File S2); to collect near Diego de Almagro in October of that year would have been inconsistent with their pace and pattern of travel. Moreover, regional mislabeling for locality names that occur in multiple regions is known for the Toro and PUCV collections (L. Ruz, personal communication) where the type material was originally held. Following these considerations, I believe the type locality to be misattributed, and have adjusted it to the village in Region IV accordingly. Diagnosis. Males are diagnosable by the combination of malar space 1.5-1.6× as long as the clypeal lateral, apical surface of the S1 process weakly longitudinally concave, and LOT above or just tangent to upper margin of anterior tentorial pits. Females are diagnosable by the combination of protibia with yellow colouration on dorsal surface only in basal half, malar space 1.4-1.6× as long as the clypeal lateral, and LOT above or just tangent to upper margin of anterior tentorial pits. Shorter-faced individuals of both sexes may be confused with C. vina and are best distinguished by the position of the LOT, which in C. vina passes through the upper margin of the anterior tentorial pits. Additionally, males of C. diaguita have the S8 anteromedial sclerotized margin wide and subparallel to margin throughout, not swollen as in C. vina. Description. Male. Length 6.0-6.7mm, forewing length 3.5-4.2mm, head width 1.2-1.4mm, thorax width 1.3-1.5mm, median ocellar diameter (OD) 0.13-0.15mm.

Chilicola (Heteroediscelis) diaguita Toro & Moldenke, 1979
Colouration: Black-brown, following parts yellow: labrum except apical margin; inverted T shape on clypeus and apex dark laterad; lower paraocular area, extending <1OD above transverse portion of epistomal suture medially, further laterally but not reaching lower tangent of antennal socket; apicoventral spot on scape; apicoventral surface of pedicel; protrochanter apical rim posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including dorsal half of anterior surface; narrow apical ring on mesofemur; narrow basal ring and apicodorsal spot on mesotibia; wide basal ring on metatibia, as well as apical quarter in ventral half only; basal one-third of metabasitarsus ventrally; faint anterior spot on tegula. Ventral surface of antennal flagellum yellow-brown, narrowing basally on F1, and F11 brown. Apical onethird of prodistitarsus brown. Metasoma black, T1-T7 marginal zones translucent yellow.
Colouration: Black to black-brown except as follows: mandible with yellow basal spot, translucent yellow in apical half except apex translucent brown. Following parts brown: pedicel, F1 and F2, on apicoventral margins. Following parts variably yellow to yellow-brown: ventral surface of F3-F10, each suffused with brown basally; dorsal surface of protibia in basal half; apical half of prodistitarsus; basidorsal spot on mesotibia; faint anterior spot on tegula. Metasoma black, T1-T5 marginal zones translucent yellow.
Material studied (

Diagnosis.
Males are separated -along with C. travesia -by the tibiae and tarsi mostly or entirely yellow-orange, and differentiated from the latter by MOC less than 0.95× width of the head, and apical surface of S1 process longitudinally concave except for a median anterior convexity. Females are separated from all other species of the subgenus except C. travesia by the combination of malar space less than 2/3× as long as the clypeal lateral, metatibia expanded (>0.25× as deep as long), and T3 with apicolateral patches of tomentum as on T1-T2. Females of C. erithropoda are differentiated from those of C. travesia by the hind tibia mostly dark (at most basal one-third and narrow apical band yellow) and the probasitarsus mostly yellow-brown to dark brown (at most basal and apical quarter yellow) -both are mostly or entirely yellow in the latter species. Description. Male. Length 5.5-6.8mm, forewing length 3.4-3.9mm, head width 1.3-1.5mm, thorax width 1.3-1.5mm, median ocellar diameter (OD) 0.13-0.15mm.
Colouration: Black-brown except as follows: antenna variable (scape with yellow apicoventral spot to entirely dark; apicoventral surface of pedicel yellow to brown; ventral surface of flagellum yellow to yellow-brown, narrowing basally on F1). Following parts yellow: labrum; clypeus except narrowly dark at mid-length along epistomal suture and apex often translucent brown laterad; lower paraocular area, extending more than 1OD above transverse portion of epistomal suture medially and reaching lower tangent of antennal socket laterally; protrochanter apical rim posteriorly; apical one-third to half of profemur on anterior surface, narrowly on posterior surface; protibia; protarsus; apex of mesofemur broadly on anterior surface, narrowly on posterior surface; mesotibia; mesobasitarsus variable from brown to yellow; apical half of mesodistitarsus; broad apical ring on metafemur; metatibia except ventral and posterior carinae brown; meta- basitarsus ventrally, sometimes dorsally in basal half; anterior spot on tegula; apex of T7. Apicoventral rim of metacoxa and metatrochanter yellow-brown. Metasoma dark brown, T1-T6 marginal zones yellow-orange to translucent yellow at margins.
Colouration: Black to black-brown except as follows: labrum entirely black or with yellow-brown apex; mandible with yellow basal spot, otherwise translucent yellow with apex translucent brown; antenna variable (scape dark or with brown apicoventral spot; apicoventral rim of pedicel yellow-brown to brown; apicoventral spot on F1 and F2 yellow-brown to brown; ventral surface of flagellum from F3 to terminal flagellomere yellow to yellow-brown). Following parts yellow: protibia except dark medioventrally; probasitarsus at most in basal and apical quarter; prodistitarsus; wide basal and apical rings on mesotibia; mesobasitarsus variable from brown to yellow ventrally; apical half of mesodistitarsus; narrow apicodorsal rim on metafemur; metatibia at most in basal one-third and narrow apical band; metabasitarsus ventrally; apical half of metadistitarsus yellow to yellow-brown; anterior spot on tegula. Metasoma black, T1-T5 marginal zones yellow to translucent yellow at margins.
Comments. The holotype is labeled Atacama, Pueblo Hundido, x.1972, L. Ruz, on Heliotropium stenophyllum. This refers to the renamed village of Diego de Almagro (Region III), but is inconsistent with the otherwise-documented distribution of C. erithropoda, as well as the known range of H. stenophyllum (Luebert 2013). Following evidence presented in the comments on C. deserticola above, I believe the type locality to be misattributed, and have adjusted it to the village of Pueblo Hundido in Region IV accordingly. Diagnosis. Males are diagnosable by the combination of malar space 1.4-2× as long as the clypeal lateral, and apical surface of S1 process with a longitudinal median ridge. Males share many characters with C. vicugna and C. mavida, but are differentiable by the longer face and weakly toothed posterior carina on the metatibia. Females are diagnosable by the combination of malar space 1.2-1.7× as long as the clypeal lateral, and dorsal surface of the protibia yellow along its entire length.
Colouration: Black-brown, following parts yellow: labrum; clypeus except narrowly dark brown along epistomal suture and apex yellow along entire width; lower paraocular area extending more than 1OD above transverse epistomal suture, nearly reaching lower tangent of antennal socket; apicoventral surface of scape; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including anterior surface in dorsal half, with ventral half translucent yellow; probasitarsus; apicodorsal rim of mesofemur, wider anteriorly; wide basal and apical rings on mesotibia, as well as apical half of anterior surface; mesobasitarsus, suffused with brown distad; apicoventral rim of metacoxa; wide apical ring on metafemur; metatibia except black posterior carina, medial dark patch on posterior surface, dark medial quadrangle on anterior surface covering approximately half of total length; basal half of metabasitarsus and ventrally in apical half; anterior spot on tegula; apex of T7. Following parts yellow-brown: apicoventral surface of pedicel; ventral surface of F1, narrowing basally; ventral surface of antenna from F2 to terminal flagellomere, suffused with darker brown; protrochanter apical rim posteriorly; apical half of prodistitarsus; apicoventral rim of mesocoxa; apical half of mesodistitarsus; apicoventral rim of metatrochanter. Metasoma black, T1-T6 marginal zones amber to translucent yellow at margins.
Colouration: Black to black-brown except as follows: labrum pale to translucent yellow along basal margin; mandible with yellow basal spot and pale to translucent yellow except apex translucent brown. Following parts yellow-brown: apicoventral surface of pedicel and F1; ventral surface of antenna from F2 to terminal flagellomere, narrowing basally on F2; mesobasitarsus ventrally; apical half of mesodistitarsus; wide apical ring on metatibia; posterior surface of metabasitarsus. Following parts yellow: dorsal surface and narrow apical ring on protibia; anterior surface of probasitarsus and second to fourth tarsomeres; apical half of prodistitarsus; wide basal and narrow apical ring on mesotibia; narrow apical ring on metafemur; wide basal ring on metatibia; anterior spot on tegula. Apical one-third of metadistitarsus brown. Metasoma brown, T1-T5 marginal zones amber to translucent yellow at margins.
Type material ( Variation. Some females have a yellow-brown or yellow spot on the lower paraocular area below the anterior tentorial pit. Distribution. Upper Intermediate Desert and northern Central Andean Cordillera, from Fundo La Semilla west to Last Juntas and north to E of Potrerillos (Region III); 875-3000m a.s.l.
Ecology. Recorded September to December and March to April.
Etymology. This species is named after the guanaco -Lama guanicoe (Müller, 1776) -a camellid native to South America and found throughout Chile. The specific epithet is identical to that of the guanaco and is treated as a noun in apposition. Diagnosis. Males are diagnosable by the combination of malar space subequal to or longer than clypeal lateral (1-1.1×), the S1 proces apical surface weakly longitudinally concave and in lateral view anterior and posterior margins divergent apically, and the metatibia >2× as long as maximum depth with summit of ventral convexity situated at approximately three-quarters tibial length. Females are diagnosable by the combination of protibia yellow on basal half of dorsal surface only and malar space subequal in length to the clypeal lateral (0.95-1.05×). Females may be confused with C. randolphi or C. vina, but can be distinguished from the former by the dark brown metasoma and from the latter by the shorter malar space (≥1.2× as long as clypeal lateral in C. vina).
Type material ( Variation. Paratype males differ in colour from the holotype as follows: inverted T shape on clypeus yellow; lower paraocular maculation extending above transverse portion of epistomal suture; wide apical ring on profemur yellow; apical half of prodistitarsus yellow-brown; narrow basal ring and large apicodorsal spot on mesotibia yellow; wide basal and apical rings on metatibia yellow, expect dark apicodorsally; basal half of metabasitarsus yellow ventrally. Additionally, some females have a yellow or yellow-brown basal spot on the mandible, the apicoventral margin of F1 and F2 brown or yellowbrown, and/or a yellow-brown or brown spot on the lower paraocular area below the anterior tentorial pit. The male S8 anteromedial sclerotized margin varies in form among paratypes: C.kate.003 has a sclerotized spot disjunct from the margin as in the holotype (easily distinguished only on the righthand lateral process), C.kate.004 has a distinct convex incursion which appears swollen, and C.kate.005 has the margin only slightly widened and weakly convex; all three were collected from Los Queñes on the same date.
Ecology. Recorded October to March and June. Etymology. The specific epithet honours Katie D'Angelo, whose support, love, and encouragement were vital to the completion of this work. It is formed in the genitive singular case.
Comments. The holotype is parasitized, with a visible female strepsipteran protruding from beneath T4. In order to verify that its morphology was not altered by parasitism, the holotype was thoroughly compared to the male paratypes for differences in colouration, pubescence, surface sculpture and structure; only colouration differed appreciably from the paratypes and these differences are noted in the variation section above. One female paratype (C.kate.007) from the same locality as the holotype is also parasitized, with two female strepsipterans protruding from beneath T4. Diagnosis. Males are diagnosable by the combination of malar space ~2.2× as long as clypeal lateral, and apical surface of S1 process with a longitudinal median ridge. Females are diagnosable by the combination of malar space ~2× as long as clypeal lateral, and the presence of wide yellow basal and apical rings on the metatibia.
Colouration: Black-brown, following parts yellow: labrum; clypeus except broadly dark brown along epistomal suture and apex yellow along entire width; lower paraocular area extending more than 1OD above transverse epistomal suture, not reaching lower tangent of antennal socket; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including dorsal half of anterior surface; apicodorsal rim of mesofemur, wider anteriorly; apicodorsal rim of metatibia; wide basal and apical rings on mesotibia, as well as apical quarter of anterior surface; basal half of mesobasitarsus dorsally; apicodorsal rim of metafemur; anterior surface of metatibia in basal quarter and apical band narrow dorsally to wide on summit of ventral convexity, posterior surface in basal one-third and apical one-third; ventral metatibial carina; basal half of metabasitarsus ventrally; apex of T7. Following parts yellow-brown: apicoventral surface of scape and pedicel, ventral surface of antenna from F1 to terminal flagellomere, narrowing basally on F1, and suffused with darker brown on remaining flagellomeres; apical half of prodistitarsus. Protrochanter apical rim posteriorly yellow-brown. Faint yellow anterior spot on tegula. Apicoventral rim of metacoxa and metatrochanter translucent brown. Posterior metatibial carina black. Metasoma black, T1-T6 marginal zones amber to translucent yellow at margins.
Colouration: Black to black-brown except as follows: labrum translucent brown with apex yellow; mandible with yellow basal spot, translucent yellow with apex translucent brown; pale spot on lower paraocular area below anterior tentorial pit. Following parts yellow: dorsal surface and narrow apical ring on protibia; probasitarsus; apical two-thirds of prodistitarsus; wide basal and apical rings on mesotibia; apical half of mesodistitarsus; apical ring on metafemur, wider ventrally; wide basal and apical ring on metatibia; anterior spot on tegula. Following parts yellow-brown: small spot on apicoventral rim of scape; apicoventral surface of pedicel, F1, and F2; ventral surface of antenna from F3 to terminal flagellomere, with pale apical bands; ventral half of metabasitarsus. Metasoma black-brown, T1-T5 marginal zones narrowly yellow to translucent yellow at margins.
Colouration: Black-brown, following parts yellow: labrum; clypeus except broadly dark along epistomal suture; lower paraocular area, extending ~1OD above transverse portion of epistomal suture and not reaching lower tangent of antennal socket; apicoventral spot on scape; apicoventral surface of pedicel; protrochanter apical rim posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; protibia except dark medioventrally; probasitarsus dorsally, at least in basal half; apex of mesofemur broadly on anterior surface, narrowly on posterior surface; broad basal and apical rings on mesotibia, the latter widened to apical half anteriorly; at least basoventral spot on mesobasitarsus; apicoventral rim of metacoxa and metatrochanter, the latter sometimes yellow-brown; narrow apical ring on metafemur; metatibia in basal one-third, in apical quarter of anterior surface, apical one-third of posterior surface and along ventral varina, yellow colouration continuous along ventral surface, sometimes continuous or nearly so along dorsal surface; metabasitarsus in basal one-third dorsally, basal two-thirds ventrally; anterior spot on tegula; apex of T7. Ventral surface of flagellum yellow-brown, flagellomeres suffused with brown basally. Apical half of prodistitarsus and apical one-third of mesodistitarsus brown. Posterior metatibial carina black. Metasoma black, T1-T6 marginal zones translucent yellow.
Colouration: Black to black-brown except as follows: mandible with yellow basal spot, translucent yellow in apical half except apex translucent brown; antenna variable (scape entirely dark; pedicel to F2 dark, or apicoventral rim brown; ventral surface of F3 to penultimate flagellomere yellow-brown, narrowing basally on F3, flagellomeres suffused with brown basally; terminal flagellomere yellow-brown in basal half, brown in apical half). Following parts yellow: dorsal surface of protibia; apical two-thirds of prodistitarsus, sometimes yellow-brown; narrow basal ring on mesotibia; apicodorsal rim on metafemur; wide basal ring on metatibia, narrow ventrally; anterior spot on tegula. Following parts yellow-brown: apicodorsal spot on mesotibia; posterior apical rim on metatibia. Apical half of mesodistitarsus brown. Metasoma black, T1-T5 yellow-brown beyond premarginal line to translucent yellow at margins.
Structure: Labrum 2.5× wider than long (20:8); malar space subequal in length to clypeal lateral (6:6.5); LOT at level of anterior tentorial pits; clypeus ~1.1× as long as maximum width in frontal view (29:26) extending for approximately half of its length Variation. Some females have a yellow-brown or yellow spot on the lower paraocular area below the anterior tentorial pit. Diagnosis. Males are diagnosable by the combination of first flagellomere longer than wide (≥1.1×), malar space shorter than clypeal lateral (<0.95×), and S1 process with anterior apical angle broadly rounded in lateral view. Females are diagnosable by the combination of clypeus strongly convex and in lateral view bulging ~1.5OD above the compound eye, metatibia expanded (≥0.25× as deep as long), and T3 apicolateral hair band sparse, not tomentose apicolaterally as on T1-T2.
Colouration: Black-brown, following parts yellow: labrum; inverted T on clypeus, with apex dark laterad; lower paraocular area extending to level of transverse epistomal suture medially, further laterally but not reaching lower tangent of antennal socket; apicoventral spot on scape and pedicel; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including dorsal half of anterior surface; apicodorsal rim of mesofemur, wider anteriorly; wide basal and narrow apical ring on mesotibia, as well as apical half of anterior surface; apicodorsal rim of metafemur; ventral surface of metatibia, as well as dorsal surface in basal quarter, narrowly yellow apically except broad around apical lamina and posterior concavity; basal half of metabasitarsus ventrally; anterior spot on tegula; apex of T7. Following parts yellow-brown: ventral surface of antenna from F1 to terminal flagellomere, suffused with darker brown basally; apical half of prodistitarsus; posterior surface of mesobasitarsus ventrally. Posterior metatibial carina black. Metasoma black, T1-T6 marginal zones amber to translucent yellow at margins.
Colouration: Black to black-brown except as follows: labrum black to yellowbrown on apex; mandible with yellow basal spot, translucent yellow with apex black. Following parts yellow-brown: apicoventral spot on F1 and F2; ventral surface of antenna from F3 to terminal flagellomere, with pale apical bands; apicodorsal rim of metafemur; basidorsal spot on metatibia. Following parts yellow: dorsal surface of protibia; apical two-thirds of prodistitarsus; basidorsal and apicodorsal spots on mesotibia. Metasoma black, T1-T5 marginal zones yellow to translucent yellow at margins.

Chilicola (Heteroediscelis) neffi
Colouration: Black-brown, following parts yellow: labrum; variable on clypeus from inverted T to median line only, apex dark laterad; lower paraocular area extending <1OD above transverse epistomal suture, not reaching ventral tangent of antennal socket; apicoventral spot on scape; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including dorsal half of anterior surface; apicodorsal rim of mesofemur, wider anteriorly; narrow basal ring and apicodorsal spot on mesotibia; apicodorsal rim of metafemur; basal quarter of metatibia, as well as apical ring narrow on anterior surface, extending to summit of ventral convexity and widened to apical quarter on posterior surface; anterior spot on tegula. Following yellow-brown: apicoventral surface of pedicel, sometimes brown; ventral surface of flagellum, narrowing basally on F1, remaining flagellomeres variably suffused with brown basally and terminal flagellomere sometimes brown; protrochanter apical rim posteriorly; anterior surface of probasitarsus; apical half of prodistitarsus. Apical one-third of mesodistitarsus brown. Posterior metatibial carina black. Metasoma dark brown, T1-T6 marginal zones yellow-brown to translucent yellow at margins. Apex of T7 yellow to translucent yellow at margin.
Colouration: Black to black-brown except as follows: mandible with yellow basal spot, translucent yellow in apical half except apex translucent brown. Following parts yellow-brown: ventral surface of antenna from F3 to terminal flagellomere, flagellomeres suffused with brown basally, terminal flagellomere brown. Following parts yellow: dorsal surface of protibia; apical half of prodistitarsus; wide basal and narrow apical ring on mesotibia; anterior spot on tegula, sometimes faint. Following parts brown: apical one-third of mesodistitarsus; apicodorsal rim of metafemur; wide basal ring on metatibia. Metasoma variable from mostly black to mostly orange-brown, at least marginal zones of T1-T5 narrowly translucent yellow; individuals with lighter metasomae have T1-T5 marginal zones more widely yellow to translucent yellow at margins, and T1-T3 with pregradular area yellow to yellow-brown, but disc always mostly brown.
Material studied (47 males & 91 females). Holotype (male): Region II, Taltal, S 25.406°, W 70.481°, 39m, 30.ix.1972, J. Neff (AMNH); Allotype (female): same data as holotype (AMNH); Region II: one paratype female, same data as holotype, on Oxalis (AMNH); one paratype female, same locality and collector as holotype, 1.x.1972 Diagnosis. Males and females of C. packeri are differentiated from all other species of the subgenus except for C. neffi by the longitudinal depressions on the paraocular area. Males are distinguished from the latter by the narrower metabasitarsus, ≥4× as long as maximum depth (≤3.9× in C. neffi). Females are distinguished by the first three metasomal segments orange-brown, with the premarginal line of T1-T3 entirely dark or with paired lateral dark spots, and disc always entirely orange-brown (in C. neffi T1-T3 are dark brown, or at most orange-brown only in marginal zone and pregradular area, with disc always dark brown in apical half ).
Colouration: Black-brown, following parts yellow: labrum; inverted T shape on clypeus, apex dark laterad; lower paraocular area extending ~1OD above transverse epistomal suture, not reaching ventral tangent of antennal socket; apicoventral surface of scape and pedicel; ventral surface of antenna from F1 to terminal flagellomere, narrowing basally on F1, remaining flagellomeres suffused with brown basally; protrochanter apical rim posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including dorsal half of anterior surface; anterior surface of probasitarsus; apical half of prodistitarsus; apicodorsal rim of mesofemur, wider anteriorly; wide basal and apical rings on mesotibia, wider on apical anterior surface; basal spot on anterior surface of mesobasitarsus; apicoventral rim of metatrochanter; apicodorsal rim of metafemur; basal quarter of metatibia, and apical ring narrow on anterior surface, extending to summit of ventral convexity and widened to apical quarter on posterior surface; basal quarter of metabasitarsus ventrally; anterior spot on tegula. Apicoventral margin of metacoxa translucent yellow. Posterior metatibial carina black. Apex of T7 yellow-brown. Metasoma dark brown, T1-T6 marginal zones yellow-brown to translucent yellow at margins.
Variation. Some females have F1-F3 more thoroughly dark, with only the apicoventral margin brown.
Distribution. Intermediate Desert and one record from Coquimban Desert, distributed coastally from Quebrada del Potrero (Region III) south to P.N. Fray Jorge (Region IV) and inland east to Travesia (Region III); 12-1613m a.s.l.
Ecology. Recorded September to November. Etymology. The specific epithet honours Laurence Packer, the author's mentor and supervisor, and collector of the holotype specimen. It is formed in the genitive singular case.
Comments. Specimens of C. packeri had previously all been identified as C. neffi due to the longitudinal depressions on the frontal area present in both species. The former was recognized following the observation that it has a distinct, non-overlapping geographic distribution relative to C. neffi, as well as a unique DNA barcode sequence, following which morphological differences were also observed. Diagnosis. Males are readily diagnosed by the distinctly truncate S1 process with apex perpendicular to axis of process in lateral view, and metabasitarsus expanded basally (~0.33× as deep as long). Among consubgeners, only C. erithropoda has the metabasitarsus expanded thus, but is readily distinguished from C. randolphi by the entirely yellow-orange tibiae and S1 process in lateral view with anterior and posterior margins divergent apically. Females are diagnosable by the red-brown metasoma and protibia yellow only on basal half of dorsal surface. Females of C. packeri, many females of C. curvapeligrosa, and some females of C. neffi also have the metasoma red-brown, but always have the protibia yellow on the entire dorsal surface.
Colouration: Black-brown, following parts yellow: labrum; clypeus except broadly dark along epistomal suture and apex brown laterad; lower paraocular area, extending dorsad to just below lower tangent of antennal socket laterally and about 1OD above transverse portion of epistomal suture medially; apicoventral spot on scape; protrochanter apical rim posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; protibia except dark medioventrally; basal half of dorsal surface of probasitarsus; wide apical ring on mesofemur; basal and apical quarters of mesotibia, dorsal surface suffused with yellow throughout; base of mesobasitarsus; apicodorsal rim of metafemur; ventral surface of metatibia, as well as dorsal surface in basal onethird and narrowly yellow apically except broad around apical lamina and posterior concavity; basal half of metabasitarsus ventrally; anterior spot on tegula; apex of T7. Following parts yellow-brown: ventral surface of antenna from apical half of pedicel to terminal flagellomere, narrowing basad on pedicel and F1; apical half of prodistitarsus; apicoventral rim of metacoxa and metatrochanter. Ventral and posterior metatibial carinae black. Metasoma dark brown, T1-T6 marginal zones amber to translucent yellow at margins.
Colouration: Black to black-brown except as follows: labrum yellow-brown; mandible with yellow basal spot, translucent yellow in apical half except apex translucent brown. Following parts brown: spot below anterior tentorial pit; median spot on clypeus just above upper margin of anterior tentorial pits; basoventral spot on scape. Following parts yellow: dorsal surface of protibia in basal half; apical two-thirds of prodistitarsus; narrow basal ring on mesotibia; faint anterior spot on tegula. Following parts yellow-brown: apicoventral surfaces of pedicel and F1; ventral surface of antenna from F2 to terminal flagellomere, flagellomeres suffused with brown basally; apical ring on protibia; apical half of mesodistitarsus; apical ring on metafemur; basal ring on metatibia. Metasoma orange-brown, graduli of T2-T4 & S2-S4 dark and widened medially to basal half of disc on S2.
Variation. Some males have the ventral margin of the metatibia dark. Females often have the gradulus of T1 dark, and usually lack the brown spots on the clypeus and below the anterior tentorial pit.
Ecology. Collected on Phacelia (12 records). Recorded December to January and March.
Etymology. The specific epithet honours the author's father, W.F. Randolph Monckton, who nurtured an interest in science and geography, and encouraged a desire for knowledge. It is formed in the genitive singular case.

Diagnosis.
Males are separated -along with C. erithropoda -by the tibiae and tarsi mostly or entirely yellow-orange, and differentiated from the latter by MOC equal to or greater than width of the head, and apical surface of S1 process longitudinally concave. Females are separated from all other species of the subgenus except C. erithropoda by the combination of malar space less than 2/3× as long as the clypeal lateral, metatibia expanded (>0.25× as deep as long), and T3 with apicolateral patches of tomentum as on T1-T2. Females of C. travesia are differentiated from those of C. erithropoda by the metatibia mostly or entirely yellow (basal one-third and apical quarter at minimum) and the probasitarsus entirely yellow (sometimes suffused with yellow-brown at midlength on posterior surface) -both are mostly dark in the latter species. Description. Male. , forewing length 3.2-3.5mm, head width 1.2-1.3mm, thorax width 1.3mm, median ocellar diameter (OD) 0.13mm.
Colouration: Black to black-brown except as follows: labrum black to translucent yellow; mandible with yellow basal spot, otherwise translucent yellow with apex translucent brown; antenna variable (scape with yellow-brown apicoventral spot to entirely dark; apicoventral surface of pedicel yellow to brown; ventral surface of flagellum yellow, either narrowing basally on F1, or with apicoventral surface of F1 brown and narrowing basally on F2). Following parts yellow: protibia, sometimes dark medioventrally; protarsus; mesotibia, at least in basal quarter ventrally, basal one-third dorsally, and apical quarter; mesotarsus, at least basitarsus ventrally and apical half of distitarsus; broad apical ring on metafemur; metatibia, at least in basal one-third and apical quarter; metabasitarsus ventrally; apical half of metadistitarsus; anterior spot on tegula. Metasoma black, T1-T5 marginal zones yellow to translucent yellow at margins.
Surface sculpture: As in male except as follows: supraclypeal area irregularly punctate (i=1-3d) densely punctate toward lateral margin (i≤d); frontal area densely punctate Diagnosis. Males are diagnosable by the combination of malar space shorter than (~0.5×) clypeal lateral, S1 process apical surface with a distinct longitudinal ridge, and in lateral view having a distinctly acute anterior apical angle, sloping strongly toward sternum posteriorly. Females are diagnosable by the combination of malar space less than half as long as clypeal lateral, clypeus sparsely punctate medially (i≥2d), and absence of longitudinal depressions on the paraocular area.
Colouration: Black-brown, following parts yellow: labrum; clypeus except broadly dark along epistomal suture, sometimes apex brown altered; lower paraocular area, extending >1OD above transverse portion of epistomal suture and not reaching lower tangent of antennal socket; apicoventral spot on scape; apicoventral surface of pedicel; protrochanter apical rim posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, as well anterior surface and basal and apical rings narrow ventrally; probasitarsus dorsally, suffused with brown ventrally; apex of mesofemur broadly on anterior surface, narrowly on posterior surface; narrow basal and apical ring on mesotibia, the latter widened to apical one-third anteriorly; anterior surface of metatibia in basal quarter and narrowly on apical margin to summit of ventral convexity, posterior surface in basal and apical one-third and along ventral carina; metabasitarsus ventrally, darker apically; anterior spot on tegula. Following parts yellowbrown: ventral surface of flagellum, flagellomeres suffused with brown basally; apical one-third of prodistitarsus; apicoventral rim of metatrochanter; apex of T7. Posterior metatibial carina black. Metasoma black, T1-T6 marginal zones translucent yellow.
Colouration: Black to black-brown except as follows: mandible yellow except apex translucent brown, sometimes trasluscent yellow in apical half; antenna variable (pedicel and F1-F2 apicoventral surface narrowly brown to broadly yellow-brown; ventral surface of F3 to penultimate flagellomere yellow, often suffused with brown basally; terminal flagellomere yellow basally, brown apically). Following parts yellow: dorsal surface of protibia; apical one-third of prodistitarsus; narrow basal and apical rings on mesotibia; wide basal ring on metatibia, narrow ventrally, as well as apical rim posteriorly; anterior spot on tegula. Following parts yellow-brown: anterior surface of probasitarsus; apicodorsal rim of metafemur. Apical one-third of mesodistitarsus brown. Metasoma black, T1-T5 yellow-brown beyond premarginal line to translucent yellow at margins.
Colouration: Black-brown, following parts yellow: labrum except apical margin; inverted T shape on clypeus, apex dark laterad; lower paraocular area at most up to level of transverse portion of epistomal suture medially, extending further laterally but not reaching lower tangent of antennal socket; apicoventral spot on scape; apicoventral surface of pedicel; protrochanter apical rim posteriorly; apex of profemur broadly on anterior surface, narrowly on posterior surface; dorsal surface of protibia, including dorsal half of anterior surface; narrow apical ring on mesofemur; narrow basal ring and apicodorsal rim of mesotibia; wide basal ring on metatibia, narrow apical ring wider ventrally; basal half to two-thirds of metabasitarsus ventrally; anterior spot on tegula. Ventral surface of antennal flagellum yellow-brown, narrowing basally on F1, F11 brown. Apical half of prodistitarsus variably yellow to yellow-brown. Ventral metatibial carina black. Metasoma black, T1-T7 marginal zones translucent yellow.
Colouration: Black to black-brown except as follows: mandible with yellow basal spot, translucent yellow in apical half except apex translucent brown. Following parts variably brown to black: pedicel, F1 and F2, on apicoventral margins; apicodorsal rim of metafemur; wide basal ring on metatibia. Following parts variably yellow to yellow-brown: ventral surface of F3-F10, suffused with brown basally; dorsal surface

Phylogenetic analysis
Analysis of 74 morphological characters (discrete and continuous) produced one most parsimonious tree with a fit of 13.3, CI = 37.4, RI = 49.7 (Fig. 50A). Most groups were found to have negative support from symmetric resampling and bootstrap analysis, meaning that these groups were more often contradicted than supported in resampling routines; groups receiving stronger support (i-ii) are discussed below.
Analysis of 658 base pairs from CO1 sequence data produced one most parsimonious tree with a fit of 42.3, CI = 51.9, RI = 61.4 (Fig. 50B). On the molecular-based tree only, C. charizard was recovered as sister to the remaining members of the subgenus, but with near-zero support; moreover, in this arrangement, its sister group is not defined by any discernible morphological character states. The remaining groups have either negative support and are not recovered in other analyses or have low, moderate, or high support and are also recovered in the subsequent analysis (discussed below). This tree agrees with the morphology-based tree with respect to the species pairs [C. neffi + C. packeri], [C. erithropoda + C. travesia], and [C. deserticola + C. mantagua]. Both trees also support the grouping together of C. vina, C. diaguita & C. katherinae and that of C. lickana, C. guanicoe & C. vicugna, albeit in different arrangements within the groups.
Combined analysis of morphological and molecular data produced a single most parsimonious tree with a fit of 28.8, CI = 45.9, RI = 55.0 (Fig. 51). This phylogeny has higher support overall than either morphological or molecular data alone, and monophly is moderately or strongly supported by resampling for a majority of the groups found. Hereafter, larger clades within this phylogeny are referred to by number for ease of reference (Fig. 51). Groups 1-3 appear at first to be problematic due to the lack of resampling support, but each is defined by at least two synapomorphies, and well supported by additional morphological characters, which are summarized below (see Suppl. material 1 for character-annotated tree, Fig. S1). Groups 3.1.1 and 3.2 (Fig. 51) and all species pairs were recovered in both the combined and molecular-only analyses, while species pairs [C. neffi + C. packeri], [C. erithropoda + C. travesia], and [C. deserticola + C. mantagua] were consistently recovered in all three analyses. The strongest contradictions to the combined analysis phylogeny come from groups (i) and (ii) in the morphology-only result. First, group (i) excludes C. randolphi relative to the combined analysis tree, and is supported by a handful of homoplasies, particularly by the S1 process with apically divergent margins in lateral view and the malar space subequal to or longer than clypeal lateral; including C. randolphi requires a reversion in both characters. Support for this grouping is otherwise low, however, while the combined analysis recovers C. randolphi as sister to [C. vina + (C. diaguita + C. katherinae)] with very high support (group 3.1.1). Second, group (ii) notably has [C. erithropoda + C. travesia] nested deep within the subgenus sister to C. guanicoe, an arrangement supported by several morphological characters; however, this species pair has many homoplasic character state reversals in this placement, and occupies a much more parsimonious position in the combined analysis tree. Thus, the combined phylogeny is presented as the preferred result.
Monophyly of the subgenus is supported by a combination of the yellow posterior margin of the female pronotal lobe, the truncate S1 process in males, the female frontal area without longitudinal striae, the male basal vein before Cu half as long as Rs, the male S2 pubescence ≥1.5OD long basally and 1.5 longer basally than apically, and the male S7 dorsal lobe with main body transverse. The species pair [C. neffi + C. packeri] is sister to the rest of the subgenus (group 1) and is defined by two synapomorphies: the longitudinal depressions on the frontal area and the broad mesoventral lobe of the male gonoforceps. The pair additionally share a male S7 with a broad ventral lobe and apodemal arm with membranous portion completely enclosed by the sclerotized margin. Only these two species have a weakly and uniformly convex apical surface of the male S1 process, but whether this character state is synapomorphic or symplesiomorphic is ambiguous, since outgroup taxa lack a truncate S1 process with a defined apical surface. Either a convex apical surface of the S1 process is the ancestral state in Heteroediscelis, or it is derived from a longitudinally concave apical surface; both arrangements require the same number of steps, although the latter requires one fewer homoplasy. Group 1 is defined by two synapomorphies: male clypeal apex yellow along its entire width, and a strap-shaped male S7 dorsal lobe. Members of this clade also have a male ventral metatibial carina which spans half the length or more of the longitudinal concavity, and the posterior carina lacks a sharp inflection at its base (this is regained in two species). Within group 1, and sister to group 2, the species pair [C. deserticola + C. mantagua] lacks any synapomorphies, and is instead grouped by a combination of several male homoplasies: the clypeus is moderately densely punctate throughout, the S1 process has a longitudinally concave apical surface, the S2 pubescence is relatively long, the apical margin of T7 is brown or black, the S7 discal apex is sinuate, and the dorsal lobe of S7 has a long basal tuft and short apical row of setae with midlength bare. Group 2 is defined by two synapomorphies: male S1 process in lateral view with margins subparallel or divergent apically and male S7 dorsal lobe with a continuous row of setae long basally and shorter apically. Within this group, the pair [C. erithropoda + C. travesia] is defined by several synapomorphies, namely by the extensively pale leg colouration in both sexes and the unique shape of both the enlarged male metatibia and male S8 lateral process. Group 3 is defined by three synapomorphies: the malar space is subequal to or longer than the clypeal lateral, the prodistitarsus is brown or black in males, and the male mesepisternum is more sparsely punctate in front of the episternal groove than behind. Additionally, the clypeus projects for at least half its length beyond the lower ocular tangent in all but three species, a character state present also in C. penai but synapomorphic within the subgenus (an elongate clypeus and/or face has numerous independent instances throughout the Xeromelissinae). This group is subdivided into groups 3.1 and 3.2.
Group 3.1 lacks synapomorphies, but is characterized by a combination of three homoplasic states, invariant in all members: the male clypeal apex is brown laterally, the male S7 dorsal lobe has a long basal tuft and short apical row of setae with midlength bare, and the mesoventral lobe of the male gonoforceps has anterior and posterior margins concave. The group is additionally supported by the sinuate male S7 discal apex. Nested within, group 3.1.1 is defined by the structure of the male posterior metatibial carina, which is sublinear and weakly defined basally, and by the colouration of the female protibia, which is yellow only in the basal half of the dorsal surface. All members of this subgroup also have a male S1 process with longitudinally concave apical surface. The group [C. vina + (C. diaguita + C. katherinae)] is united by six invariant homoplasies: the male clypeal maculation forms an inverted T shape, the male lower paraocular maculation extends no more than 1OD above the transverse epistomal suture, the genal beard is relatively sparse, the male S8 lateral process is ≥1.2 wider than long, the apicolateral hair patches of the female terga are tomentose on T1-T2 only, and the female legs are extensively dark. The pair [C. diaguita + C. katherinae] is not defined by any readily identified morphological characters; support for this pairing comes from molecular evidence, as the two share a synapomorphic substitution of C at site 595 of CO1 (relative to Apis mellifera L.) and homoplasic substitution of A at site 531 (although two divergent sequences from C. vina have these same substitutions). Group 3.2 is defined by three synapomomorphies: the male ventral metatibial carina is absent basally, the posterior carina extends the full length of the longitudinal concavity and is weakly sigmoid, and the apical surface of the male S1 process has a distinct longitudinal median ridge in all species. As well, the male S7 dorsal lobe is triangular, a convergent character state present in C. penai but synapomorphic within Heteroediscelis. [C. lickana + (C. guanicoe + C. vicugna)] has synapomorphic female leg colouration, and is further supported by the male S1 process in lateral view with margins convergent apically and by the relatively long male S2 pubescence. The pair [C. guanicoe + C. vicugna] is united by the lower paraocular maculation reaching the antennal socket (converged upon in other species, but unique in group 3.2). [C. mayu + (C. curvapeligrosa + C. mavida)] is supported by synapomorphic male hindleg colouration, and the pair [C. curvapeligrosa + C. mavida] share a male third flagellomere subequal in length to the second (unique in group 3.2). Morrone's (2015) biogeographic areas are outlined in Fig. 52, and species distributions are depicted in Fig. 49. Analysis with VIP identified 12 disjunct sister pairs in the consensus reconstruction, with disjunctions present at 12 out of 16 internal nodes; these are indicated on Fig. 51. Each pair of sister species exhibits a north-south disjunction -i.e. along a latitudinal gradient -while divergence along an elevational gradient is evident at two nodes: first, the disjunction between group 3 and [C. travesia + C. erithropoda] lies longitudinally between the Andes and lower interior to the east, and along the boundary between the Intermediate and Coquimban Deserts to the south, thus separating dry, northern, coastal and interior areas inhabited by [C. travesia + C. erithropoda], from more humid, southern, and Andean areas inhabited by group 3. The second is the disjunction between C. charizard and group 3.1.1, which corresponds well to the boundary between the Intermediate and Coquimban Deserts at higher elevations, and at lower elevations runs north-by-northwest along coastal mountains. A third disjunction, between C. vina and [C. diaguita + C. katherinae], also lies in part along an elevational gradient, but is discussed below.

Biogeography
Of the latitudinal disjunctions, seven correspond to previously identified boundaries between biogeographic areas, and three cut across biogeographic areas, seeming to lie along divisions between river systems. Of the latter three, two are associated with the Río Huasco: the disjunction between C. vicugna and C. guanicoe crosses the Intermediate Desert somewhere between the Huasco and Copiapó river systems, while the disjunction between C. travesia and C. erithropoda lies along or just to the south of the Río Huasco. The third, a disjunction between C. mayu and [C. curvapeligrosa + C. mavida], extends west across the Santiagan province (at ~35°S) somewhere between the Mantaquito-Teno-Lontué and Maule river systems.
Out of the seven remaining disjunctions mentioned above, at least three correspond to the boundary between the Coquimban and Santiagan provinces (~32.5-33°S), giving it particularly good support; these are the disjunctions between C. katherinae and C. diaguita, between C. mavida and C. curvapeligrosa, and between the two sister clades of group 3.2. This last disjunction precedes the one between C. mavida and C. curvapeligrosa, and actually extends north along the Andes to accommodate the range of C. mavida; this can be interpreted as a disjunction at 32.5-33°S between the ranges of the northern clade [C. lickana + (C. guanicoe + C. vicugna)] and the southern clade [C. lickana + (C. guanicoe + C. vicugna)], followed by later allopatric dispersal or vicariance of C. mavida along the Andes to the north. The Coquimban-Santiagan boundary separates semi-arid scrubland to the north from more humid sclerophyll and spiny (espinal) woodland to the south, though most of this woodland has been destroyed by heavy cultivation (Chester 2008, Peña 1966, Morrone 2015. A similar disjunction is present between C. mantagua and C. deserticola, with the distribution of C. mantagua extending a little further north along the coast than the other Santiagan groups, to south of the Río Choapa. The disjunction between C. neffi and C. packeri suggests a barrier somewhat north of Copiapó (~26.5°S) near the boundary between Peña's (1966) Northern Coast and Intermediate Desert regions. The southern limit of the Northern Coast is pushed north in Morrone's regionalisation (2015), but a boundary near ~26.5°S can be defined based on moisture: to the north the coastal zone is very dry and receives nearly all of its moisture from fog, while to the south rainfall is more common (Peña 1966). Moreover, just inland, this boundary lies along the southern extent of the Interior Desert district, sometimes known as the "Absolute Desert" (Chester 2008). The most  (2015) and districts are those proposed therein, with boundaries reproduced from their respective authors (Peña 1966, O'Brien 1971, Artigas 1975. Areas marked with an asterisk (*) represent undescribed portions of the Puna province, and are given placeholder names for ease of reference. Scale bar approximately 200 km, background imagery provided by ESRI.
pronounced disjunction with regard to moisture is that between C. lickana and [C. guanicoe + C. vicugna], lying somewhere between the Coquimban province of the Central Chilean sub-region and the Puna province of the South American transition zone. Chilicola lickana's distribution is the most arid of the subgeneric range, and this species may well have diverged as a result of one or more episodes of hyperaridity of the Atacama Desert (Amundson et al. 2012, Jordan et al. 2014. The disjunction between C. vina and [C. diaguita + C. katherinae] consists of two disjunctions: one in the north which crosses the Río Elqui near Vicuña, separating higher-elevation-inhabiting C. vina from C. diaguita, and one in the south across the Andes at around 34°S, south of which C. katherinae occurs. The southern disjunction corresponds well to Peña's boundary between the Central and Southern Andean Cordillera regions near the Río Tinguiririca, which also forms the northern boundary of Peña's precordilleran Northern Valdivian Forest region (both were shifted in Morrone's regionalisation, to the north and south, respectively). To the south of this boundary, an increase in humidity is marked by the appearance of Nothofagus forests (Peña 1966, Chester 2008. Depending on which of the phylogenetic hypotheses is considered, there are at least two possible interpretations for this apparent two-fold disjunction. Under the preferred phylogeny, one interpretation is that these three species diverged from one another in a relatively short time, which could help explain the discrepancy between molecular and morphological evidence as to whether C. vina or C. katherinae diverged first. Alternatively, in the morphology-only phylogeny, C. katherinae is sister to the other two species (Fig. 50A); if this is taken as the true relationship between these three species, it would suggest two separate disjunctions, with the southern one preceding the northern one.

Taxonomy
This revision increases the size of the subgenus Heteroediscelis; with the description of eight new species and the redescription of nine previously-described ones, the total is increased to seventeen (previously ten). Also included are three minor, but notable alterations to the taxonomic record: the synonymy of C. mantagua with C. valparaiso, the designation of a neotype specimen for C. vicugna, and a re-interpretation of the type localities of C. erithropoda and C. deserticola. Details regarding the latter two are included under comments on the relevant species descriptions. The first was motivated by insufficient distinction between C. mantagua and C. valparaiso. The diagnosis for C. valparaiso refers to two characters : malar space subequal to minimum width of scape, and apical lamina of metatibia narrowing distally in apical view. In fact, the holotypes of C. mantagua and C. valparaiso both have the malar space approximately half as long as the clypeal lateral (i.e. of equal relative length), and the apical lamina of the metatibia is not differentiable between them. Moreover, characters of the male terminalia do not support any division between these two supposed species, and I have identified many specimens from the same broad geographic area as the C. valparaiso holotype as either C. mantagua or C. vina. Indeed, the type locality of C. valparaiso is just to the south of the originally-documented range of C. mantagua. Finally, CO1 barcode sequences from specimens tentatively identified as C. valparaiso are indistinguishable from those of C. mantagua. Therefore, in absence of any evidence to maintain the distinction of C. valparaiso as a species, I have treated it as a synonym of C. mantagua -the latter is the type species of the subgenus and has page priority in the original publication .

DNA Barcodes
DNA barcoding was useful for the association of females to males of several species, particularly among those whose females are difficult to tell apart (e.g. C. deserticola, C. erithropoda, C. mantagua, and C. vicugna). As an exploratory tool, barcoding initially helped reveal the presence of two new species from localities new to the subgenus: C. randolphi from Laguna del Maule and C. curvapeligrosa from Farellones. Barcode sequences also served as supporting evidence for the description of morphologically distinct species, such as C. charizard, C. lickana, and C. mayu. In three cases, however, DNA barcodes of distinct species clustered into a single BIN. These are: C. mavida & C. curvapeligrosa, C. vicugna & C. guanicoe, and C. vina, C. diaguita & C. katherinae ( Table 1). All seven of these species are supported by the morphological characters provided in the keys and descriptions, but are reduced to three when using BINs as a proxy for species (i.e. molecular operational taxonomic units -MOTUs; Floyd et al. 2002). On the other hand, there are three species which are represented by seven total BINs: C. mantagua (two BINs), C. neffi (three BINs) and C. packeri (two BINs); as a result, BIN analysis overestimates the number of MOTUs found at low elevations and underestimates the number of MOTUs found at high elevations. DNA barcoding is indeed a rapid, efficient means to estimate diversity (e.g. Smith et al. 2005, Gibson et al. 2014) and can be invaluable as an aid in the discovery of new species (e.g. Gibbs 2009a, 2009b, Packer et al. 2009, Butcher et al. 2012), but in this specific case, at least, using BINs to evaluate patterns of diversity seems not to produce accurate results.
As a partial correction, a higher resolution of molecular species identification is possible by specifying diagnostic, single-nucleotide substitutions within barcode sequences (Fisher and Smith 2008, Gibbs 2009a, 2009b. I have summarized these for Heteroediscelis in Table 1. Of the seventeen species in the subgenus, thirteen have diagnostic nucleotide subsitutions (or diagnostic combinations of them) in their barcode sequences. Provided a specimen belongs to the subgenus, the presence of any one of these substitutions should result in succesful identification of the corresponding species. In particular, this approach allows molecular differentiation of C. mavida and C. curvapeligrosa, as well as separation of C. katherinae from C. diaguita & C. vina; it also allows multiple BINs associated with a single species to be collapsed into a single MOTU. Given that diagnostic nucleotide subsitutions are very easily determined, the practice of identifying them in taxonomic revisions should be encouraged (Fisher and Smith 2008, Gibbs 2009a, 2009b.

Phylogeny and biogeography
The phylogeny in Fig. 51 represents the most strongly supported evolutionary hypothesis for the subgenus, drawing on a total evidence approach to analysis (Eernisse and Kluge 1993, Huelsenbeck et al. 1996. A single tree was recovered with good GC/BS support for most of its branches, and with ample morphological support for all of them. Aside from the morphological evidence supporting the various clades, this phylogeny includes groupings which have some biogeographic basis. Disjunctions in distributions of sister clades correspond in many cases to the biogeographic areas described by Morrone (2015) and Peña (1966), and in a few cases to boundaries identified by others. For Table 1. Diagnostic or invariant DNA barcode nucleotide substitutions, and Barcode Index Numbers (BINs) for each species of Chilicola (Heteroediscelis). Nucleotide position is given relative to the CO1 sequence of Apis mellifera. Brackets indicate subtitutions which must co-occur in order to be diagnostic. Nucleotides in other species are listed in order of prevalence. BINs which are not unique to one species are italicized.

Diagnostic nucleotides
Most common nucleotide in other species Barcode Index Number (BIN) C. mantagua 5 115 -C 640 -C example, both the southern disjunction between C. vina and [C. diaguita + C. katherinae] and the disjunction between C. mayu and [C. curvapeligrosa + C. mavida] correspond to a vicariance event identified by Dominguez et al. (2016) in tenebrionid beetles and phasmids, and indeed Peña's boundary between the Central and Southern Andean Cordillera falls in the same general area, representing the northern limit of moderate-elevation Nothofagus woodlands (Peña 1966, Chester 2008. As well, the disjunction between C. lickana and [C. guanicoe + C. vicugna] unambiguously places these taxa on either side of an occasionally-cited summer/winter rainfall divide across northern Chile, which arose 2-2.2 Mya (Amundson et al. 2012), marking the boundary between the tropical and mid-latitude climate zones (Arroyo et al. 1988, Messerli et al. 1993, Houston 2006. While a few of the earlier divergences have associated disjunctions across an elevational gradient, most of the recent divergences exhibit disjunctions across the latitudinal moisture gradient which extends from the arid north to the temperate south. These disjunctions may be the result of varying degrees of aridification in northern Chile, given that the Atacama Desert reached its present hyperarid state sometime after the origin of Chilicola , Dunai et al. 2005, Rech et al. 2010, and perhaps around the same time as the origin of Heteroediscelis (Jordan et al. 2014). In any case, divergence along this gradient is the primary pattern for more closely related taxa, and in many cases coincides with well-documented shifts in vegetation, such as the shift from semi-arid scrubland to sclerophyll and spiny woodland across the Coquimban-Santiagan boundary, and the appearance of Nothofagus woodlands south of ~34°S noted above (Peña 1966, Chester 2008. Although ancestral distributions cannot be inferred from the analysis used in the present study, one can speculate: the grade comprising [C. neffi + C. packeri], [C. deserticola + C. mantagua], and [C. erithropoda + C. travesia] is collectively distributed along the primarily coastal and low-elevation portions of the Coquimban & Intermediate Deserts, which could suggest an ancestral distribution along the semi-arid mid-northern coast and interior (commonly called the Norte Chico; Sánchez and Morales 1990). This BOLD:AAI4251 † this BIN includes one sequence from C. vina, from which C. guanicoe and C. vicugna can be separated by the indicated substitution. * this BIN contains only one sequence from C. lickana and as such the substitutions in quotation marks may not be diagnostic; a second, partial (284 bp) sequence from C. lickana shares the first three substitutions, but does not span the remaining three bases. would be consistent with the designation of Heteroediscelis as endemic to Chile, with Andean (i.e. high-elevation) and latitudinally more extreme distributions potentially representing derived ranges.
With the completion of this phylogenetic analysis, in addition to the work done by Gibbs and Packer (2006) on Chilicola s. str. and by Willis and Packer (2007) on Chilioediscelis, revisionary work on the clade [Heteroediscelis + (Chilicola s. str. + Chilioediscelis)] found in Packer's subgeneric phylogeny (2008) is fit for synthesis. Combined, these three revisions provide an exhaustive set of data well-suited to a thorough analysis of the clade, which could incorporate discrete morphological, continuous morphological, and molecular evidence. In light of this -along with the eight species newly described herein, plus additional new discoveries in Chilicola s. str. and Chilioediscelis (L. Packer, personal communication) -a combined phylogenetic analysis of these three subgenera together with Oediscelis is recommended in order to re-assess subgeneric boundaries and relationships within Chilicola, and to investigate biogeographic patterns throughout its broad neotropical distribution.

Data matrix
Morphological data used in the phylogenetic analysis is summarized in the tables below. For DNA barcode data used in the phylogenetic analysis and for the specification of diagnostic nucleotide substitutions, see Suppl. material 3: DNA Barcode Data. Table A1. Discrete morphological data used in phylogenetic analysis.