Corresponding author: Spencer K. Monckton (
Academic editor: M. Ohl
The bee subgenus
Monckton SK (2016) A revision of
The subgenus
A number of new species belonging to
For the revision, 1316 specimens of Packer Collection at Pontificia Universidad Católica de Valparaíso, Chile Bee Biology and Systematics Laboratory, Logan, UT, U.S.A. Central Texas Melittological Institute
Specimens were identified using the existing key (
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 Canadian Centre for DNA Barcoding Barcode of Life Data Systems database Barcode Index Numbers
Accordingly, barcode sequences were used in species delimitation employing an integrative approach (e.g.
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
Specimen records not originally having GPS coordinates are supplemented with them where possible, as determined via discussions with the collectors and using
The following acronyms are used: : diameter of the median ocellus : distance from lower tangent of median ocellus to apex of clypeus : ocellocular distance, shortest distance between lateral ocellus and compound eye : interocellar distance, shortest distance between lateral ocelli : upper & lower interocular distance, shortest distance between inner orbits of compound eyes, above & below level of emargination, respectively : maximum interocular distance, or greatest distance between inner orbits of compound eyes at level of emargination : lower ocular tangent, an imaginary line crossing lower extremities of compound eyes in frontal view, perpendicular to main axis of face : interalveolar distance, shortest distance between antennal sockets : minimum antennocular distance, shortest distance between antennal socket and compound eye, typically at an angle of approximately 30° below horizontal laterad from antennal socket : antennal flagellomere : tergum : sternum
Descriptions use terminology from
Distributions were plotted using ArcGIS 10.3.1 (
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.
Outgroups were selected based on the findings of
Specimens were examined for morphological characters that were potentially phylogenetically informative, and additional characters were adapted from the key and diagnoses of
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;
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
Representative
Maximum parsimony analysis was performed using extended implied weighting ( consistency index retention index bootstrap
Biogeographic areas referenced throughout follow Morrone’s formal regionalisation (
Distributions of : Vicariance Inference Program
Males are readily diagnosable by S1 with a robust, elongate, ventrally directed process, with truncate apex having a well-defined (sometimes oblique) apical surface (other
Small bees, length 4.2–6.8mm.
Known from Chile, as far north as San Pedro de Atacama (Region II) and south to Cuesta Las Raíces (Region IX). Distributed throughout the Central Chilean sub-region, and found also in the Puna province (South American transition zone) and Maule province (Subantarctic sub-region); 0–3210m a.s.l.
Habitats range from
Note: High magnification is required for some identifications; an ocular micrometer capable of distinguishing differences of 0.01mm is recommended. In frontal view, the perimeter of the median ocellus and the apex of the clypeus must be in the same plane of focus.
1 | Tibiae and tarsi mostly or entirely yellow-orange (Fig. |
|
– | Tibiae and tarsi mostly black-brown (Fig. |
|
2(1) | Apical surface of S1 process longitudinally concave throughout (Fig. |
|
– | Apical surface of S1 process concave posteriorly, convex anteromedially (Fig. |
|
3(1) | Frontal area with longitudinal depression above each antennal socket which partially accommodates the scape (Fig. |
|
– | Frontal area without such depressions (Fig. |
|
4(3) | Metabasitarsus ≤3.8× as long as maximum depth (Fig. |
|
– | Metabasitarsus ≥4× as long as maximum depth (Fig. |
|
5(3) | Apical surface of S1 process with distinct longitudinal median ridge, surface otherwise approximately flat (Fig. |
|
– | Apical surface of S1 process either longitudinally concave (Fig. |
|
6(5) | Malar space ≥1.4× as long as clypeal lateral (Fig. |
|
– | Malar space ≤1.1× as long as clypeal lateral (Fig. |
|
7(6) | Clypeus with a distinct median longitudinal depression (Fig. |
|
– | Clypeus with median longitudinal depression weak (Fig. |
|
8(6) | Malar space ~0.5× as long as clypeal lateral (Fig. |
|
– | Malar space ≥0.8× as long as clypeal lateral (Fig. |
|
9(8) | Malar space longer than (1.05–1.1×) clypeal lateral (Fig. |
|
– | Malar space subequal to or shorter than (0.8–1×) clypeal lateral (Fig. |
|
10(8) | First flagellomere subequal in length and width or shorter (<1.05×) than wide (Fig. |
|
– | First flagellomere ≥1.1× as long as wide (Fig. |
|
11(5) | Malar space longer than (>1.05×) clypeal lateral (Fig. |
|
– | Malar space shorter than (<0.95×) clypeal lateral (Fig. |
|
12(11) | Malar space very long, ≥2.7× as long as clypeal lateral (Fig. |
|
– | Malar space not as long, ≤1.6× as long as clypeal lateral (Fig. |
|
13(12) | Metatibia broadly expanded (≥0.5× as deep as long) and with summit of ventral convexity near to tibial apex, at approximately 0.85× tibial length (Fig. |
|
– | Metatibia less expanded (>2× as long as maximum depth) and with summit of ventral convexity situated basad of tibial apex at approximately three-quarters of tibial length (Fig. |
|
14(13) | Malar space 1.5–1.6× as long as clypeal lateral (Fig. |
|
– | Malar space 1.2–1.4× as long as clypeal lateral (Fig. |
|
15(11) | In lateral view, truncate apex of S1 process perpendicular to axis of process, anterior and posterior margins parallel apically (Fig. |
|
– | In lateral view, truncate apex of S1 process slightly oblique, anterior and posterior margins convergent apically (Fig. |
|
16(15) | Mesobasitarsus mostly yellow (Fig. |
|
– | Mesobasitarsus mostly dark (Fig. |
|
1 | Protibia with yellow on dorsal surface present in basal half only, apical half dark (Fig. |
|
– | Protibia with yellow along entire dorsal surface (Fig. |
|
2(1) | Malar space at most 0.8× as long as clypeal lateral (Fig. |
|
– | Malar space at least as long as clypeal lateral (Fig. |
|
3(2) | In frontal view, lower ocular tangent passing just tangent to or above upper margin of anterior tentorial pits (Fig. |
|
– | In frontal view, lower ocular tangent passing through anterior tentorial pits (Fig. |
|
4(3) | Malar space ≤1.1× as long as clypeal lateral (Fig. |
|
– | Malar space ≥1.2× as long as clypeal lateral (Fig. |
|
5(1) | Malar space clearly longer than (>1.1×) clypeal lateral (Fig. |
|
– | Malar space at most subequal in length to clypeal lateral (Fig. |
|
6(5) | Malar space less than <1.7× as long as clypeal lateral (Fig. |
|
– | Malar space at least twice as long as clypeal lateral (Fig. |
|
7(6) | Metatibia with wide yellow basal and apical rings (Fig. |
|
– | Metatibia entirely dark, or with a wide basal band of brown that is clearly darker than the pale colouration on other legs (Fig. |
|
8(5) | Malar space approximately 0.9× as long as, or subequal in length to, clypeal lateral (Fig. |
|
– | Malar space at most two-thirds as long as clypeal lateral (Fig. |
|
9(8) | Apex of metafemur and base of metatibia yellow-brown to brown, distinctly darker than same parts of midleg (Fig. |
|
– | Apex of metafemur and base of metatibia yellow, as on same parts of midleg (Fig. |
|
10(8) | Frontal area with longitudinal depressions which partially accommodate the scape (Fig. |
|
– | Frontal area without such depressions (Fig. |
|
11(10) | Metasoma with at least T1–T3 orange-brown, except for dark premarginal line and/or a pair of lateral dark spots (Fig. |
|
– | Metasoma darker, at most T1–T3 orange-brown with apical half of disc dark (Fig. |
|
12(10) | T3 with apicolateral hair band sparse, not tomentose as on T1–T2 (Fig. |
|
– | T3 with apicolateral patches of tomentum as on T1–T2 (Fig. |
|
13(12) | Clypeus strongly convex in dorsal half, bulging ~2OD above compound eye in lateral view (Fig. |
|
– | Clypeus not as above, dorsal half bulging ≤ |
|
14(12) | Metatibia somewhat expanded, >0.25× as deep as long (Fig. |
|
– | Metatibia not expanded, and >4.1× as long as maximum depth (Fig. |
|
15(14) | Metatibia mostly yellow, at least in basal one-third and apical quarter, often entirely so (Fig. |
|
– | Metatibia mostly dark, at most basal one-third and a narrow yellow apical ring (Fig. |
|
16(14) | Metapostnotum with sparse, irregular longitudinal carinae (Fig. |
|
– | Metapostnotum rugose, often more weakly so posteriorly (Fig. |
|
Male, leg colouration:
Male:
Male, frontal view:
Male:
Male S1, apical surface:
Male:
Male, frontal view:
Male:
Male:
Male:
Male:
Male:
Male:
Male:
Male:
Male:
Female, dorsal surface of protibia:
Female:
Female, frontal view:
Female:
Female:
Female:
Female:
Female:
Female, frontal view:
Female:
Female:
Female:
Female:
Female:
Female:
Males are diagnosable by the combination of malar space short (~0.6× clypeal lateral), mesobasitarsus mostly dark, and metatibia with the combination of ventral carina extending full length of longitudinal concavity and smooth, 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 (~0.5× clypeal lateral), metapostnotum with sparse, irregular longitudinal carinae, and protibia yellow along entire dorsal surface but darkened somewhat to yellow-brown apically. Among consubgeners
(38 males & 20 females):
Coastal, southern Coquimban Desert, from Peñuelas (Region V) north to Pichidangui (Region IV); 13–362m a.s.l.
Collected on
This species is diagnosable by the malar space, which is the longest in the subgenus: in males it is ~2.7× as long as the clypeal lateral, in females it is ~2.33× as long as the clypeal lateral.
(6 males & 11 females):
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.
Intermediate Desert & Central Andean Cordillera, from Fundo La Semilla (Region III) south to Elqui Prov., 6 km S of Vicuña (Region IV) and west to Incahuasi & Embalse Sta. Juana (Region III); 645–2358m a.s.l.
Collected on
The specific epithet is in homage to the fictional monster which this species resembles. It is treated as a noun in apposition.
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
(27 males & 49 females).
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
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.
Collected on
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.
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).
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). Among consubgeners,
(58 males & 110 females).
Coquimban & Intermediate Desert, distributed coastally from Caleta Chañaral de Aceituna (Region III) south to Los Vilos (Region IV) and inland along the Elqui Valley to Pueblo Hundido & Guanta (Region IV); 8–1691m a.s.l.
Collected on
The type series (male holotype & one male paratype) is labeled Atacama, Pueblo Hundido,
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
(8 males & 22 females):
Coquimban–Intermediate Desert transition zone, from Choros Bajos southeast to Tololo Observatory (Region IV); 20–1686m a.s.l.
Recorded September to November and February to March.
Males are separated - along with
(19 males & 22 females):
Coastal, southern Intermediate Desert, from Chañaral de Aceituna (Region III) south to Los Choros (Region IV), with type locality disjunct from otherwise observed distribution; 8–1031m a.s.l.
Collected on
The holotype is labeled Atacama, Pueblo Hundido,
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
(9 males & 26 females):
Some females have a yellow-brown or yellow spot on the lower paraocular area below the anterior tentorial pit.
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.
Recorded September to December and March to April.
This species is named after the guanaco -
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
(5 males & 17 females):
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 yellow-brown, 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.
Southern Andean Cordillera, from Río Teno near Estero Pichuante west to Los Queñes (Region VII) and south to Valle Las Trancas (Region VIII); 707–1549m a.s.l.
Recorded October to March and June.
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.
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.
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.
(3 males & 3 females).
Northern Volcanic Cordillera, from Aguas Blancas north to 12 km N of San Pedro de Atacama (Region II); 2474–2801m a.s.l.
Recorded October and December.
The specific epithet is taken from Kunza, an extinct language of the Atacama people.
Males are diagnosable by the combination of first flagellomere longer than wide (≥1.1×), malar space longer than clypeal lateral (1.05–1.1×), and S1 process with anterior apical angle acute in lateral view. Females are diagnosable by the combination of malar space subequal in length to the clypeal lateral (0.9–1×) and leg colouration as follows: apicodorsal spot on mesotibia yellow-brown, apicodorsal rim of metafemur yellow, wide basal ring of metatibia yellow, posterior apical rim of metatibia yellow-brown. Females of
(21 males & 19 females):
Some females have a yellow-brown or yellow spot on the lower paraocular area below the anterior tentorial pit.
Central Andean Cordillera and upper Coquimban Desert, just extending into Southern Andean Cordillera, from Baños del Toro & Embalse La Laguna west to El Pangue (Region IV) and south to Portillo (Region V); 1142–3210m a.s.l.
Collected on
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.
(5 males & 7 females).
Pehuén and Southern Andean Cordillera, from Cuesta Las Raíces (Region IX) north to Laguna del Maule (Region VII) and west to Valle Las Trancas (Region VIII); 1221–2227m a.s.l.
Collected on
The specific epithet was recommended by R.A. Lühr and is derived from the type locality near Lonquimay, a place name formed in part from the Quechua
Males and females of
(47 males & 91 females).
Northern coastal extension of Intermediate Desert, from Taltal north to 29 km N of Paposo and inland SE of Taltal (Region II); 5–676m a.s.l.
Collected on
One female from Ruta 5, km 1006 (C.neff.072) has the metasomal colouration of
Males and females of
(42 males & 96 females).
Some females have F1-F3 more thoroughly dark, with only the apicoventral margin brown.
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.
Recorded September to November.
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.
Specimens of
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
(6 males & 15 females):
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.
Southern Andean Cordillera, from Laguna del Maule (Region VII) southwest to Valle Las Trancas (Region VIII); 783–2541m a.s.l.
Collected on
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.
Males are separated - along with
(29 males & 65 females):
Intermediate Desert, distributed coastally from Balneario Flamenco south to Huasco Bajo and inland to Travesía (Region III); 0–713m a.s.l.
Collected on
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.
(84 males & 201 females).
Some females have a yellow-brown or yellow spot on the lower paraocular area below the anterior tentorial pit.
Central Andean Cordillera and Coquimban & Intermediate Desert, from Embalse Sta. Juana (Region III) south to Las Mollacas (Region IV), west to El Tofo, and east to E of Vicuña, Hwy. 41-CH km 137.8 (Region IV); 406–1991m a.s.l.
Collected on
Designation of the neotype is justified because of damaged type material; identification of
Males are diagnosable by the combination of malar space 1.2–1.4× as long as the clypeal lateral, apical surface of the S1 process weakly longitudinally concave, and the
(58 males & 71 females):
Coquimban Desert, Central Coastal Cordillera, Central Valley, and Southern Andean Cordillera, from Viña del Mar (Region V) north to Pisco Elqui (Region IV), south to Alto de Cantillana (RM Santiago), and east to E of El Volcán (RM Santiago); 15–1963m a.s.l.
Distribution of
Collected on
Analysis of 74 morphological characters (discrete and continuous) produced one most parsimonious tree with a fit of 13.3,
Results of maximum parsimony analysis of:
Analysis of 658 base pairs from
Combined analysis of morphological and molecular data produced a single most parsimonious tree with a fit of 28.8,
Single most parsimonious tree resulting from maximum parsimony analysis of combined morphological and molecular dataset. GC support values are indicated above branches, bootstrap support values are below. Nodes with distributional disjunctions are indicated by a black square. Numbered groups are discussed in text.
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 [
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 [
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 [
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
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 [
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
Chilean biogeographic areas overlaid with distribution of
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
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
The disjunction between
The disjunction between
This revision increases the size of the subgenus
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.
Diagnostic or invariant DNA barcode nucleotide substitutions, and Barcode Index Numbers
Species | Number of sequences | Diagnostic nucleotides | Most common nucleotide in other species | Barcode Index Number (BIN) |
---|---|---|---|---|
|
5 | 115 - C |
A |
|
|
5 | 19 - A |
T,C |
|
|
5 | 232 - A |
A,T |
|
|
5 | 241 - T | A |
|
3 |
- | - |
|
|
|
6 | 223 - C |
T,A |
|
6 |
337 - C | T |
|
|
|
3 | 133 - A |
A |
|
|
1 (+1)* | 125 - A |
G |
|
|
4 | 232 - T |
A,T |
|
|
2 | 364 - C |
T
|
|
|
2 | 124 - A | T | |
|
4 | 292 - G |
A |
|
|
4 | 5 - C |
T
|
|
|
11 | 124 - C |
T
|
|
this BIN includes one sequence from
this BIN contains only one sequence from
As a partial correction, a higher resolution of molecular species identification is possible by specifying diagnostic, single-nucleotide substitutions within barcode sequences (
The phylogeny in Fig.
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
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
Although ancestral distributions cannot be inferred from the analysis used in the present study, one can speculate: the grade comprising [
With the completion of this phylogenetic analysis, in addition to the work done by
I express my warmest thanks to Dr. Laurence Packer for his mentorship, support, and enthusiasm. His patience, understanding, and genuine care are endlessly appreciated. To Katie, for her incredible support, love, and encouragement: I love you. I thank my family for their care and attention: Mum for her love and understanding, Dad for his continued interest and encouragement, my siblings for their enthusiasm, and the D’Angelos for their engagement and support. Additional thanks to those who assisted with fieldwork in Chile: to Alfredo Ugarte, Rolando Humire Coca, and Roberto Lühr for their help and hospitality, and to Rosie Smith for her help and accompaniment. Special thanks to James Postlethwaite, an excellent field companion, for his friendship. I am grateful also to the members of the Packer Lab, in particular: Tom Onuferko, J. Scott MacIvor, Sheila Dumesh. Thanks to volunteers Shammi and Shane for help recording specimen data. This work would not have been possible without specimens from the
Characters 1–54 are represented by discrete states; characters 55–74 are continuous and based upon ratios.
Male, colour of clypeal apex: (0) yellow full width (Fig.
Male, clypeus colour: (0) entirely yellow except along epistomal suture (Fig.
Male, clypeal median depression: (0) absent or marked by a very weak/faint depression (Fig.
Male, clypeal sculpture: (0) moderately densely punctate (i=1-2d) throughout; (1) sparsely punctate (i=2-3d) medially.
Male, F1: (0) at most as long as wide (Fig.
Male, ratio of length of F2:F1: (0) subequal, <1.1x (Fig.
Male, ratio of length of F3:F2: (0) subequal, <1.05x; (1) ≥1.1x.
Male, frontal line: (0) flat/indistinct, or carinate only between antennal bases; (1) carinate for 1/2 to 3/4 distance from supraclypeal area to median ocellus.
Male, lower paraocular maculation: (0) extending <1OD above transverse portion of epistomal suture (Fig.
Female, colour of F1 and F2 ventrally: (0) mostly brown or black (Fig.
Female, clypeal sculpture: (0) moderately densely punctate throughout, i=1-2d (Fig.
Female, ratio of length of F3:F2: (0) subequal, ≤1.1×; (1) ≥1.2×.
Female, longitudinal striae on frontal area: (0) absent (
Genal beard: (0) moderately dense in male and sparse in female (Fig.
Projection of clypeus: (0) projecting for less than half its length beyond
Clypeus above anterior tentorial pit: (0) unmodified, projecting ≤1OD above compound eye in lateral view (Figs
Frontal area: (0) without longitudinal depressions (Figs
Malar space: (0) <0.9× clypeal lateral (Fig.
Male, metatibia shape: (0) 2.0-2.2× as long as maximum depth, summit of ventral convexity at two-thirds to three-quarters tibial length (Fig.
Male, ventral metatibial carina: (0) greatly reduced, originating in apical half of posterior carina (Fig.
Male, posterior metatibial carina: (0) absent basally, originating in apical half of ventral carina (Fig.
Male, midleg colour: NRLC axis 1 (See Appendix
Male, midleg colour: NRLC axis 2 (See Appendix
Male, hindleg colour: NRLC axis 1 (See Appendix
Male, hindleg colour: NRLC axis 2 (See Appendix
Male, mesepisternum: (0) punctures anterior of episternal groove same density as those below scrobe; (1) punctures anterior of episternal groove sparser than those below scrobe.
Male, prodistitarsus: (0) entirely brown or black; (1) apical one-third/half yellow-brown or yellow.
Male, basal vein length before Cu: (0) at most half as long as Rs (
Female, pronotal lobe colour: (0) dark; (1) with yellow posterior margin.
Female, mesepisternum: (0) punctures anterior of episternal groove same density as those below scrobe; (1) punctures anterior of episternal groove sparser than those below scrobe.
Female, prodistitarsus: (0) entirely brown or black; (1) apical one-third/half yellow-brown or yellow.
Female, protibia dorsal surface: (0) yellow for entire length (Fig.
Female, mid-/hindleg colour: NRLC axis 1 (Appendix
Female, mid-/hindleg colour: NRLC axis 2 (Appendix
Metatibial spurs: (0) unsclerotised and straight (
Metepisternum: (0) without longitudinal striae anterodorsally; (1) distinctly longitudinally striate anterodorsally.
Male, S1: (0) slightly swollen apicoventrally (Packer 2007: Fig.
Male, S1 process apical surface: (0) longitudinally concave (Fig.
Male, S1 process anterior and posterior margins in profile: (0) convergent apically (Fig.
Male, S2 pubescence: (0) ≥1.5OD long basally and 3× longer basally than apically; (1) ≥1.5OD long basally and 1.5× longer basally than apically; (2) reduced, no longer than 1OD basally.
Male, T7 apical margin: (0) brown or black; (1) yellow.
Male, S7 ventral lobe: (0) narrow, almost linear (Fig.
Male, S7 dorsal lobe orientation: (0) main body transverse, basal portion recurved anterolaterally (
Male, S7 dorsal lobe shape: (0) roughly triangular, wide at base narrowing to apex (Fig.
Male, S7 dorsal lobe setation: (0) row of setae long basally and shorter apically (Fig.
Male, S7 apex of disc: (0) sinuate (32H); (1) distinctly emarginated (Fig.
Male, S7 apodemal arm: (0) sclerotized margin entire, enclosing membranous portion of arm (Fig.
Male, S8 lateral process: (0) anteromedial sclerotized margin following marginal contour (Fig.
Male, S8 lateral process: (0) ratio of width:length ~1-1.1×, apicolateral margin evenly rounded or weakly concave (Fig.
Male, S8 spiculum: (0) short (
Male, gonobase, apicoventral truncate process: (0) biconvex and roundly emarginated (Fig.
Male, gonoforceps, mesoventral lobe: (0) evenly rounded (Fig.
Female, metasoma: (0) mostly black-brown (Fig.
Female, terga: (0) T1-T3 with apicolateral patches of tomentum (Figs
Male, ratio of clypeus width:minimum distance between subantennal sutures.
Female, ratio of clypeus width:minimum distance between subantennal sutures.
Male, ratio of subantennal suture length:minimum distance between them.
Female, ratio of subantennal suture length:minimum distance between them.
Male, ratio of
Female, ratio of
Male, ratio of scape length:maximum width.
Male, ratio of UOD:LOD.
Male, ratio of
Female, ratio of
Male, ratio of probasitarsus length:depth.
Female, ratio of probasitarsus length:depth.
Male, ratio of prodistitarsus length:combined length of tarsomeres 2-4.
Female, ratio of prodistitarsus length:combined length of tarsomeres 2-4.
Male, ratio of metafemur length:depth.
Female, ratio of metafemur length:depth.
Female, ratio of metatibia length:depth.
Male, ratio of metabasitarsus length:depth.
Male, ratio of metapostnotum length:mesoscutum length.
Female, ratio of metapostnotum length:mesoscutum length.
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
Discrete morphological data used in phylogenetic analysis.
Character # | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | 24 | 25 | 26 | 27 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
|
1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | ? | 3 | 1 | 2 | 0 | 2 | 0 | 1 |
|
1 | 0 | 2 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 2 | 0 | 0 | - | - | - | 3 | 3 | 3 | 2 | 0 | 1 |
1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 2 | 0 | 0 | 0 | 0 | 2 | 0 | ? | 1 | 2 | 1 | 0 | 0 | 1 | |
|
1 | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 3 | 0 | 2 | 0 | 0 | 2 | 0 | 2 | 1 | 0 |
|
0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 2 | 1 | 2 | 2 | 2 | 2 | 2 | 1 | 0 |
|
0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 2 | 0 | 3 | 3 | 1 | 3 | 0 | 1 |
|
1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 2 | 2 | 3 | 1 | 0 | 2 | 0 | 2 | 1 | 0 |
|
0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 2 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 3 | 0 | 5 | 2 | 5 | 2 | 0 | 1 |
|
0 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 2 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 2 | 0 | 1 | 3 | 3 | 2 | 4 | 2 | 1 | 0 |
|
1 | 1 | 2 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 2 | 0 | 0 | 2 | 0 | 2 | 1 | 1 |
|
0 | 0 | 2 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 3 | 0 | 1 | 3 | 3 | 1 | 1 | 1 | 0 | 0 |
|
0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 1 | 2 | 1 | 2 | 0 | 1 |
|
0 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 2 | 1 | 2 | 3 | 2 | 3 | 2 | 1 | 0 |
|
1 | 1 | 2 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 2 | 0 | 2 | 0 | 0 | 2 | 1 | 2 | 3 | 1 | 2 | 2 | 1 | 0 |
|
1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 3 | 0 | 2 | 1 | 2 | 0 | 1 |
|
1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 3 | 3 | 0 | 1 | 2 | 0 | 1 |
|
1 | 0 | 2 | 1 | 0 | 1 | 0 | 1 | 2 | 1 | 1 | 1 | 0 | 2 | 0 | 1 | 0 | 0 | 2 | 3 | 1 | 4 | 2 | 3 | 2 | 1 | 0 |
|
0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 2 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 3 | 0 | 5 | 2 | 5 | 2 | 0 | 1 |
|
0 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 2 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | 3 | 0 | 1 | 0 | 1 | 1 |
|
1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 2 | 2 | 3 | 1 | 0 | 2 | 0 | 2 | 1 | 0 |
Character # | 28 | 29 | 30 | 31 | 32 | 33 | 34 | 35 | 36 | 37 | 38 | 39 | 40 | 41 | 42 | 43 | 44 | 45 | 46 | 47 | 48 | 49 | 50 | 51 | 52 | 53 | 54 |
|
1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | - | - | 2 | 0 | 1 | 0 | 2 | 3 | 0 | 1 | 0 | 2 | 0 | 1 | 0 | 0 | 1 |
|
0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | - | - | 2 | 1 | 2 | 2 | 0 | 2 | 0 | 0 | 0 | 2 | 1 | 2 | 0 | 0 | 0 |
0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 1 | - | 0 | 2 | 1 | 4 | 1 | 2 | 1 | 2 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | |
|
1 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 2 | 1 | 2 | 1 | 0 | 0 | 0 | 1 | 2 | 0 | 2 | 0 | 0 | 1 | 1 | 1 | 0 | 0 |
|
1 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 1 | 2 | 3 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 1 |
|
1 | 1 | 1 | 0 | 0 | 3 | 0 | 0 | 1 | 2 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | 0 | 1 | 0 | 1 | 1 | 2 | 0 | 0 | 0 |
|
1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 2 | 0 | 2 | 1 | 0 | 0 | 0 | 1 | 2 | 0 | 1 | 0 | 2 | 1 | 1 | 1 | 0 | 1 |
|
1 | 1 | 0 | 1 | 0 | 5 | 0 | 0 | 0 | 2 | 1 | 2 | 2 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 3 | 1 | 2 | 1 | 0 | 0 |
|
1 | 1 | 1 | 1 | 0 | 4 | 0 | 0 | 0 | 2 | 3 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 2 | 0 | 0 | 0 |
|
1 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 2 | 0 | 2 | 1 | 0 | 1 | 0 | 1 | 2 | 1 | 2 | 2 | 2 | 1 | 2 | 1 | 0 | 1 |
|
1 | 1 | 0 | 1 | 0 | 4 | 0 | 0 | 1 | 2 | 3 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 |
|
1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 2 | 0 | 0 | 0 | 0 | 3 | 0 | 1 | 2 | 0 | 0 | 0 | 0 | 1 | 2 | 0 | 0 | 1 |
|
1 | 1 | 1 | 0 | 0 | 1 | 2 | 0 | 1 | 2 | 3 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 2 | 0 | 0 | 0 |
|
1 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 1 | 2 | 3 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 2 | 2 | 0 | 2 | 1 | 0 | 1 | 0 | 1 |
|
1 | 1 | 1 | 1 | 0 | 2 | 0 | 0 | 1 | 2 | 2 | 0 | 1 | 0 | 3 | 0 | 2 | 1 | 1 | 0 | 1 | 1 | 1 | 2 | 2 | 0 | 1 |
|
1 | 1 | 1 | 0 | 0 | 2 | 0 | 0 | 0 | 2 | 2 | 0 | 1 | 1 | 3 | 0 | 2 | 1 | 1 | 0 | 0 | 1 | 1 | 2 | 2 | 1 | 1 |
|
1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 2 | 0 | 1 | 1 | 1 | 3 | 0 | 1 | 2 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 |
|
1 | 1 | 0 | 1 | 0 | 6 | 0 | 0 | 0 | 2 | 0 | 2 | 2 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 2 | 3 | 1 | 1 | 0 | 0 | 0 |
|
1 | 1 | 1 | 0 | 0 | 4 | 0 | 0 | 1 | 2 | 3 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 2 | 0 | 0 | 1 |
|
1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 2 | 0 | 2 | 0 | 0 | 2 | 0 | 1 | 2 | 0 | 1 | 1 | 2 | 1 | 1 | 1 | 0 | 1 |
Continuous morphological data used in phylogenetic analysis.
Character # | 55 | 56 | 57 | 58 | 59 | 60 | 61 | 62 | 63 | 64 |
---|---|---|---|---|---|---|---|---|---|---|
|
0.523 | 0.317 | 0.492 | 0.459 | 0.220 | 0.000 | 0.804 | 1.000 | 1.000 | 1.000 |
|
0.176 | 0.063 | 0.056 | 0.000 | 0.000 | 0.361 | 1.000 | 0.000 | 0.573 | 0.788 |
0.294 | 0.133 | 0.000 | 0.170 | 0.047 | 0.265 | 0.727 | 0.528 | 0.569 | 0.604 | |
|
0.075 | 0.744 | 0.761 | 1.000 | 0.805 | 0.638 | 0.359 | 0.822 | 0.205 | 0.139 |
|
0.150 | 0.194 | 0.340 | 0.477 | 0.715 | 0.587 | 0.283 | 0.585 | 0.296 | 0.357 |
|
0.411 | 0.333 | 0.391 | 0.419 | 0.337 | 0.806 | 0.172 | 0.700 | 0.296 | 0.540 |
|
0.411 | 0.151 | 0.726 | 0.541 | 0.285 | 0.451 | 0.144 | 0.621 | 0.231 | 0.185 |
|
0.929 | 0.641 | 0.590 | 0.442 | 0.166 | 0.587 | 0.364 | 0.238 | 0.098 | 0.099 |
|
0.689 | 0.556 | 0.596 | 0.649 | 0.805 | 0.491 | 0.127 | 0.700 | 0.296 | 0.185 |
|
0.541 | 0.052 | 0.449 | 0.435 | 0.185 | 0.357 | 0.227 | 0.443 | 0.320 | 0.250 |
|
1.000 | 1.000 | 1.000 | 0.911 | 0.439 | 0.331 | 0.144 | 0.310 | 0.296 | 0.117 |
|
0.495 | 0.296 | 0.391 | 0.326 | 0.220 | 0.683 | 0.283 | 0.506 | 0.350 | 0.409 |
|
0.579 | 0.333 | 0.619 | 0.387 | 1.000 | 0.491 | 0.127 | 0.630 | 0.220 | 0.359 |
|
0.144 | 0.342 | 0.333 | 0.360 | 0.461 | 0.477 | 0.327 | 0.581 | 0.410 | 0.439 |
|
0.882 | 0.744 | 0.695 | 0.567 | 0.303 | 1.000 | 0.394 | 0.519 | 0.325 | 0.326 |
|
0.156 | 0.444 | 0.275 | 0.485 | 0.387 | 0.587 | 0.080 | 0.432 | 0.189 | 0.200 |
|
0.012 | 0.226 | 0.181 | 0.287 | 0.659 | 0.740 | 0.000 | 0.310 | 0.536 | 0.339 |
|
0.901 | 0.280 | 0.488 | 0.329 | 0.366 | 0.587 | 0.127 | 0.524 | 0.000 | 0.000 |
|
0.000 | 0.000 | 0.272 | 0.288 | 1.000 | 0.806 | 0.727 | 0.906 | 0.350 | 0.295 |
|
0.147 | 0.171 | 0.434 | 0.414 | 0.303 | 0.331 | 0.227 | 0.822 | 0.296 | 0.270 |
Character # | 65 | 66 | 67 | 68 | 69 | 70 | 71 | 72 | 73 | 74 |
|
1.000 | 0.972 | 0.952 | 0.228 | 0.463 | 0.929 | 1.000 | 1.000 | 0.000 | 0.000 |
|
0.219 | 0.341 | 0.674 | 0.528 | 1.000 | 0.000 | 0.000 | ? | 0.471 | 0.990 |
0.054 | 0.000 | 0.806 | 0.528 | 0.665 | 0.240 | 0.443 | 0.419 | 0.559 | 0.516 | |
|
0.470 | 1.000 | 0.000 | 0.041 | 0.859 | 0.844 | 0.512 | 0.527 | 0.658 | 0.761 |
|
0.235 | 0.273 | 0.525 | 0.817 | 0.350 | 0.532 | 0.548 | 0.313 | 0.919 | 0.993 |
|
0.425 | 0.451 | 0.226 | 0.272 | 0.457 | 0.562 | 0.656 | 0.185 | 0.680 | 0.813 |
|
0.387 | 0.574 | 0.348 | 0.325 | 0.658 | 0.740 | 0.561 | 0.548 | 0.698 | 0.711 |
|
0.381 | 0.304 | 0.195 | 0.281 | 0.000 | 0.395 | 0.377 | 0.000 | 0.680 | 0.300 |
|
0.378 | 0.439 | 0.613 | 0.249 | 0.855 | 0.740 | 0.634 | 0.536 | 0.587 | 0.757 |
|
0.473 | 0.295 | 0.392 | 0.427 | 0.866 | 0.694 | 0.532 | 0.916 | 0.722 | 0.717 |
|
0.357 | 0.709 | 0.645 | 0.528 | 0.482 | 0.565 | 0.781 | 0.512 | 0.813 | 0.662 |
|
0.219 | 0.273 | 0.419 | 0.469 | 0.857 | 0.428 | 0.548 | 0.185 | 0.503 | 0.501 |
|
0.193 | 0.345 | 0.636 | 1.000 | 0.708 | 0.629 | 0.689 | 0.263 | 0.613 | 0.751 |
|
0.098 | 0.169 | 0.525 | 0.240 | 0.280 | 0.642 | 0.392 | 0.244 | 0.839 | 0.923 |
|
0.271 | 0.473 | 0.613 | 0.672 | 0.399 | 0.680 | 0.522 | 0.341 | 0.758 | 0.489 |
|
0.425 | 0.451 | 0.015 | 0.336 | 0.717 | 0.740 | 0.586 | 0.441 | 0.513 | 1.000 |
|
0.000 | 0.439 | 1.000 | 0.413 | 0.625 | 1.000 | 0.728 | 0.042 | 1.000 | 0.969 |
|
0.425 | 0.473 | 0.000 | 0.000 | 0.263 | 0.795 | 0.313 | 0.325 | 0.566 | 0.359 |
|
0.219 | 0.615 | 0.226 | 0.208 | 0.412 | 0.502 | 0.598 | 0.542 | 0.786 | 0.610 |
|
0.425 | 0.709 | 0.636 | 0.312 | 0.866 | 0.844 | 0.542 | 0.726 | 0.993 | 0.703 |
Three character state trees were constructed for non-redundant linear coding of potentially confounded leg colour characters (
Male, midleg colour (original characters: apex of mesofemur, base of mesotibia, midlength of mesotibia, apex of mesotibia).
Male, hindleg colour (original characters: apex of metafemur, base of metatibia, midlength of metatibia, apex of metatibia).
Female, mid-/hindleg colour (original characters: apex of mesofemur, base of mesotibia, apex of mesotibia, apex of metafemur, base of metatibia, apex of metatibia).
Character-Annotated Phylogeny
Phylogeny
Figure S1. Phylogeny with evolutionary character changes mapped onto branches. Character numbers are above branches with corresponding states below. Solid circles indicate unique changes, open circles indicate homoplasies. Character optimization is colour-coded as black (unambiguous), red (fast), or blue (slow).
Material Studied
Occurrence & collection data
Specimen data for 1316 specimens, including localities, geographic coordinates, and depositories.
DNA Barcode Data
Sequence data