Research Article |
Corresponding author: Remko Leijs ( remko.leijs@samuseum.sa.gov.au ) Academic editor: Michael Ohl
© 2017 Remko Leijs, Michael Batley, Katja Hogendoorn.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Leijs R, Batley M, Hogendoorn K (2017) The genus Amegilla (Hymenoptera, Apidae, Anthophorini) in Australia: A revision of the subgenera Notomegilla and Zonamegilla. ZooKeys 653: 79-140. https://doi.org/10.3897/zookeys.653.11177
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The Australian bees in the subgenera Notomegilla and Zonamegilla of the genus Amegilla are revised. Commonly in Australia the species in these subgenera are called blue-banded bees, although not all species have blue bands. A phylogeny based on mitochondrial cytochrome oxidase 1 sequence data was used to delineate the species and a set of morphological criteria was developed for species identification. Strong support was obtained for separating the Australian species into the three subgenera previously proposed on the basis of morphology. Two species, are recognised in the subgenus Notomegilla and eleven new synonymies are proposed. Twelve Australian species are recognised in the subgenus Zonamegilla including four new species: indistincta, karlba, paeninsulae and viridicingulata, and twenty new synonymies are proposed. Keys to the species of both sexes and descriptions or redescriptions of all species are provided. Distribution maps, data on flower visitation and phenology are given.
blue-banded bee, pollinator, taxonomic revision, mtDNA phylogeny
Many species in the genus Amegilla are conspicuous because of their colourful, iridescent hair bands, relatively large size and hovering flight patterns. Some species are known to nest in sand, loam or clay soils, and in soft sandstone, clay washouts and mud bricks. Nests are often found in large aggregations that are re-used for many generations (
The life-cycles of the different species seem to have many similarities. With a life-span of approximately 6 weeks, adult Amegilla are relatively short-lived. Depending on the species and the suitability of the climate, one or several generations are produced during a flight season, while immatures survive the unfavourable season in the prepupal stage in their cells (
The taxonomy of the genus Amegilla has had a long and colourful history. Amegilla zonata (Linnaeus, 1758) was among the first organisms to receive a Latin binomial. The first Australian bee species to be described, Amegilla cingulata (Fabricius, 1775) was collected by Sir Joseph Banks.
Prior to 1940, species confusion and name changes were frequent, due in no small part to morphological similarities between species. In addition, many morphological character traits of Amegilla seemed variable, both within and between species, which made it difficult to find reliable characters to distinguish the species. Conscious of this intraspecific variation,
Here we present a revision of the species in the subgenera Notomegilla and Zonamegilla based on examination of the majority of the type material and supported by molecular phylogenetics based on mitochondrial DNA of all currently recognised Australian species sampled throughout their geographical ranges.
This study is based on examination of museum specimens. The following acronyms are used in the supplementary information associated with this paper:
AMS Australian Museum, Sydney
AQIS Australian Quarantine and Inspection Service, Cairns
ASCU Orange Agricultural Institute, Agricultural Scientific Collections Unit
BMNH The Natural History Museum, London
MVMA Museum of Victoria Entomology, Melbourne
WINC Waite Insect and Nematode Collection, Adelaide University, Waite Campus, Adelaide
Information from the labels of museum specimens was copied in a single database, which was subsequently used to generate distribution maps, analyse species phenology and flower records. The database is available as supporting information associated with this publication (Suppl. material
Fresh specimens were collected throughout Australia for molecular analysis by the authors and by numerous other people acknowledged below. Special attention was given to type localities for named species and localities known from apparently undescribed species in museum collections. Some non-Australian A. (Zonamegilla) species were included to enable inferences about the historical biogeography of this subgenus. Collected specimens were killed and preserved in absolute ethanol to allow DNA extraction at a later stage. Ethanol-preserved specimens, as well as extracted DNA, are kept in the Australian Biological Tissue Collection at the South Australian Museum. DNA voucher specimens are kept in the Entomology Collection of the South Australian Museum. Locality data, voucher numbers and GenBank Accession numbers of specimens used in the molecular analyses are available as supporting data associated with this publication (Suppl. material
Genitalia were extracted from a number of male museum specimens, after relaxation for up to three days in a humid container with chromocresol or thymol added as a fungal inhibitor. Dissected genitalia were treated with 10% cold sodium hydroxide for 24 hours, glacial acetic acid for approximately one hour and stored in glycerol to facilitate the study of morphology. Digital photography using an automontage system was used to obtain images of relevant characters. Morphological terminology follows that of
Characters of S7 of male Zonamegilla species, ventral view. The broadly oval head at the posterior end of S7 has a medial excavation in the dorsal surface laterally and a longitudinal elevation or ridge on the ventral surface. The excavation is carinate laterally and the sclerotisation between the carinae and the ventral ridge is usually thinner than the surrounding areas creating windows in the pigmentation. Most of the medial ridge is covered with short hairs which become longer and denser towards the middle of the head, creating a T- or Y-shaped brush. a apodeme b ventral ridge c posterior margin d medial ridge e apical projection f lateral carina of dorsal excavation g neck.
The description of, and differentiation between, intra- and interspecific colour variation has for a long time bedevilled the taxonomy of this group of bees. Colour variation in both the iridescent hairs of the tergal bands and the non-reflective hairs of the mesosoma are found in most species and can, in some cases, obscure genuine intraspecific colour differences. The metallic interference colours, which are produced by reflection from arrays of microtubules (
To overcome the difficulty of separating interspecific from intraspecific morphological variation, we used partial mitochondrial CO1 gene sequences, as in the ‘barcoding’ approach (
DNA extraction, PCR amplification and sequencing were performed as described in
Phylogenetic analyses of aligned sequence data were carried out using the program PAUP* version 4.0b8 (
A BEAST analysis allowed the application of relaxed molecular clock methods in order to obtain estimates of node divergence times. Because fossils are not known for Amegilla bees a mean rate of 0.0115 substitutions per site per million years (
MRBAYES analysis was performed with four simultaneous chains, sampling tree topologies and parameters every 50 generations. After 2,029,200 generations all parameters had reached their ESS, and the potential scale reduction parameter was approximately one for all parameters, indicating that the Bayesian runs had converged and that a sufficient sample of the posterior distribution had been obtained. A burn in of 10,000 sampled trees was chosen for each independent run of MRBAYES. A >50% posterior probability consensus tree was constructed from the remaining 61,170 saved trees (30,585 trees per individual run).
Phylogenetic analyses of CO1 gene sequences, using neighbour-joining and parsimony in PAUP* and Bayesian methods in MRBAYES and BEAST resulted in very similar tree topologies with three monophyletic groups (Fig.
Morphological and molecular evidence indicates the existence of only two valid species in Notomegilla, and twelve valid Australian species in Zonamegilla. With two exceptions the uncorrected pairwise distances between specimens resulted in unambiguous delineation of the species with maximum intraspecific distances mostly less than 4.5% and interspecific distances greater than 5.8% (Table
The second exception involved difficulties with delineation within the species-group that includes A. pulchra (Smith), A. murrayensis (Rayment) and A. adelaidae (Cockerell). In all analyses, the group as a whole was well defined with adelaidae as sister to the other two species. Within the group, initially analyses of sequences obtained using the M202/M70 primers resulted in two clades for murrayensis and a single clade with rather large intraspecific divergences for specimens of pulchra (Suppl. material
Maximum percentage uncorrected pairwise intraspecific sequence divergence of Australian Amegilla and distances to nearest other species clade A) based on 822 bp fragment of CO1 using primers M202/M70, B) based on 648 bp amplified using primers M414/M423. ‘-‘: no data available. ‘*’: inflated pairwise divergence due two multiple clades possible resulting from nuclear paralogues. (See also supplementary Figs S1 and S2)
Subgenus | % maximum intraspecific divergence A |
% distance to nearest other species clade A |
n | % maximum intraspecific divergence B |
% distance to nearest other species clade B |
n | |
---|---|---|---|---|---|---|---|
Notomegilla | aeruginosa | 0.4 | 17.2 | 7 | - | - | - |
chlorocyanea | 2.6 | 15.5 | 39 | - | - | - | |
Zonamegilla | adelaidae | 2.5 | 7.2 | 8 | 0.3 | 6.6 | 2 |
asserta | 3.7 | 7.2 | 35 | 2.2 | 5.8 | 5 | |
alpha | - | - | - | 0.2 | 6.8 | 2 | |
thorogoodi | 0.5 | 0.6 | 4 | 1.3 | - | 12 | |
cingulata | 0.5 | 6.8 | 13 | 0.0 | 6.5 | 2 | |
indistincta | 0.5 | 7.2 | 3 | 0.3 | 6.8 | 2 | |
murrayensis | 9.4* | 9.8* | 56 | 1.3 | 5.3 | 19 | |
karlba | - | - | - | - | 7.5 | 1 | |
paeninsulae | - | - | - | 3.9 | 6.7 | 4 | |
pulchra | 9.3* | 9.8* | 15 | 8.6* | 5.3 | 13 | |
walkeri | 0.8 | 0.6 | 7 | - | - | - | |
viridicingulata | 4.5 | 9.0 | 4 | - | 7.3 | 1 |
Bayesian phylogenetic analysis (Fig.
One well-supported species-group within Zonamegilla, which includes cingulata (Fabricius), paeninsulae sp. n., mcnamarae (Cockerell) from Papua New Guinea and a species from Thailand, comprises species with similar morphological characteristics including brightly coloured hair bands on the posterior margins of the terga and almost uniformly coloured hairs on the scopa of the hind tibia of the females. Among the Australian species, only viridicingulata sp. n. shares this combination of characters, but it is present in a number of Asian species. The species asserta (Cockerell) is sister to the cingulata group.
Another well-supported clade that includes A. alpha (Cockerell), thorogoodi, indistincta sp. n., karlba sp. n. and walkeri consists of species that are superficially rather different in appearance. Most notable is the orange-brown hair covering most of the metasoma in alpha, which led Cockerell to suggest that it was a subspecies of A. (Asaropoda) bombiformis (Smith). A third species-group, adelaidae, murrayensis and pulchra, contains species with partially overlapping distributions (Figs
The most useful diagnostic characters that emerged from this revision were the length of the basitibial streak on the hind leg of females, the size, shape and colour of the medial patch of pubescence on the female T5, and the shape of the male S7. The colour of the scutal pubescence, the tergal hair bands and the pubescence on female T5, although sufficiently reliable characters in fresh specimens, are less reliable in older or worn specimens. Similarly, the shape of the yellow face markings, especially in males, although often a useful indicator, can be aberrant in a small number of individuals. Characters of the male genital capsule have not been used in this study, because obvious differences between species were not evident.
Climate seems to determine the phenology of the species. Those species that have a wide latitudinal distribution (A. chlorocyanea (Cockerell), A. asserta (Cockerell) and murrayensis) are active in summer in the south of the continent, but are found year-round in the north. Species that occur only in the north of the country are most active in May and June. Along the eastern seaboards of south Queensland and New South Wales, activity peaks in October and May.
Floral records were available for a total of 583 specimens, by far the most for A. chlorocyanea (n = 447), followed by A. aeruginosa (Smith) (n = 37), murrayensis (n = 36) and thorogoodi (n = 21). The number of records available for other species ranges from 1 to 17. The bees visited 147 plant species of 47 different families (Fig.
The phylogeny estimated using BEAST (Fig.
The Australian species within the subgenus Zonamegilla are probably not monophyletic. This is indicated by species from India, Thailand and Papua New Guinea occurring amongst the Australian species (Fig.
In all but three cases, Rayment described his new species without reference to a type specimen, but listed a number of dates, localities and collectors from which a series of presumed syntypes could potentially be identified. He also left a considerable number of specimens with handwritten labels containing a species name and the word “type” or “allotype”. Only a fraction of the material referred to in Rayment’s descriptions was located and in many cases the dates given in the published information differed from those on the specimen labels (though this is not necessary for a specimen to be considered as a syntype). Although these circumstances made it tempting to treat all Rayment’s names as nomina dubia, presumed syntypes were identified, primarily from specimens with identification labels in Rayment’s distinctive hand and collection dates before February 1941, the latest date recorded in Rayment’s manuscripts. For type series containing more than one species a lectotype was chosen, though the published descriptions were of little assistance in deciding which species should be selected. The consequences of these decisions were minimal as all but two of Rayment’s names were synonymised. For each species the decision for synonymy is discussed in detail in the systematic section below.
The specimens from examined museum collections as well as newly collected material have been data-based, and were used to generate distribution maps (Figs
The whereabouts of type material for the following species is unknown or is in such condition that they could not be distinguished from other species by morphology or the original description.
Anthophora adelaidae ernesti Rayment
Anthophora adelaidae ernesti Rayment, 1947, p. 47.
Amegilla ernesti (Rayment), Michener, 1965, p. 216.
No type material bearing this name was located. Although the name was synonymised with adelaidae (Brooks, 1988) no justification was given.
Anthophora berylae Rayment
Anthophora berylae Rayment, 1947, p. 49.
Amegilla berylae (Rayment), Michener, 1965, p. 216.
Amegilla (Zonamegilla) berylae (Rayment), Brooks, 1988, p. 511.
Type material for berylae was not found.
Anthophora hackeri Rayment
Anthophora hackeri Rayment, 1947, p.55.
Amegilla hackeri (Rayment), Michener, 1965, p. 216.
Amegilla (Zonamegilla) hackeri (Rayment), Brooks, 1988, p. 511.
Syntypes of hackeri: male, Mossman, Queensland, 5 May 1940, “Type”,
While the specimens were undoubtedly the syntypes from which the species was described, they could not be distinguished from other species as the hidden sterna and genital capsule of the male are missing and the hair patch on T5 of the female is badly worn. The specimens may eventually prove to be conspecific with thorogoodi or indisctincta if ancient DNA sequence methods can be applied.
Amegilla (Zonamegilla) zonata (Linnaeus)
Apis zonata Linnaeus, 1758, p. 576.
This name has frequently been applied to Australian species and has appeared in checklists long after
Anthophora zonata cincta Sichel
Anthophora zonata cincta Sichel, 1869, p. 58.
No type material associated with this name was located. Smith (1879) pointed out that Sichel had incorrectly applied the name cincta to an Australian species and proposed emmendata as a replacement name.
Note: The following key is based on the species re-described in this paper and the majority of the specimens examined were in good condition. Because the key unavoidably includes references to colour and hair patterns that may be affected by age, wear or intraspecific variation, it should be used in conjuction with the remarks that accompany the detailed descriptions.
Females
1 | Forewing: hairs in 1st medial cell and most other cells; metasomal terga T1–4 of most species with apical hair bands | 2 |
– | Forewing: hairs absent in 1st medial cell, in other cells hairs absent or restricted to radial, marginal, 1st and 2nd submarginal cells; metasomal terga T1–4 without apical hair bands | subgenus Asaropoda |
2 | Integument of paraocular areas black; fore and mid femora and tibiae with iridescent blue-green hairs (subgenus Notomegilla) | 3 |
– | Integument of paraocular areas partly yellow, white or ivory; hair on fore and mid legs never iridescent (subgenus Zonamegilla) | 4 |
3 | Metasoma uniformly covered with green/bronze hair | aeruginosa |
– | Metasoma with pale blue bands, often with orange tints | chlorocyanea |
4 | Scutal hair various but not bright orange; hind tibial scopa white with dark longitudinal streak below basitibial plate | 5 |
– | Scutal hair orange; hind tibial scopa orange with at most very short brown mark below basitibial plate | 12 |
5 | Dark streak on hind tibial scopa < 0.5× length tibia | 6 |
– | Dark streak on hind tibial scopa ≥ 0.5× length tibia | 9 |
6 | Thoracic hair appears grey due to mixed black and pale hairs; metasomal hair bands metallic blue | walkeri |
– | Thoracic hair appears orange or pale brown; metasomal hair bands various | 7 |
7 | Metasomal hair bands matt orange, relatively wide (band on T2 about 0.4× width of disc); T5 with pale hair across full width (Fig. |
adelaidae |
– | Metasomal hair bands iridescent, but becoming dull with age; T5 with pale hair medially | 8 |
8 | Metasomal hair bands blue; dark scopal streak 0.3–0.5× length hind tibia; T5 with broad band of pale hair bordering fimbria (Fig. |
thorogoodi |
– | Metasomal hair bands yellowish; dark scopal streak 0.2–0.4× length hind tibia; T5 with medial patch of pale hair (Fig. |
indistincta |
9 | Metasomal hair bands matt orange, relatively wide (band T2 about 0.4× width of disc); T5 with pale hair across full width (Fig. |
adelaidae |
– | Metasomal bands iridescent aqua, not wide (band T2 about 0.3× width of disc); T5 with pale hair scattered and sometimes with a central line | 10 |
11 | Face marks yellow; T5 with scattered white hair and dense central line extending into prepygdial fimbria (Fig. |
asserta |
– | Face marks ivory or pale yellow; T5 with scattered white hair but central line, if present, not extending into prepygdial fimbria | 11 |
12 | Pale hair pattern on T5 wide, narrower laterally (Fig. |
murrayensis |
– | White hair pattern on T5 usually weaker, frequently with only scattered white hairs near lateral margins (Fig. |
pulchra |
13 | Metasomal terga with bands of dense orange, scale-like hair on apical margins, simpler orange hair distributed openly elsewhere | alpha |
– | Metasomal terga T1–T4 with pale hair only in apical bands | 13 |
13 | T5 medially with black hair only, some white hair at lateral margins (Fig. |
cingulata |
– | T5 with pale hair medially | 14 |
14 | Metasomal hair bands metallic orange with green iridescence; prepygidial fimbria orange-brown; T5 with scattered orange hair and medial line of denser orange hair extending into fimbria (Fig. |
paeninsulae |
– | Metasomal hair bands green, greenish-yellow or dull orange; T5 with broad patch of dispersed white hair (Fig. |
15 |
15 | Hind tibial scopa bright orange with at most slight darkening below basitibial plate; more than half hind basitarsus covered with orange hair | viridicingulata |
– | Tibial scopa usually pale orange, with a short brown streak below basitibial plate; less than half hind basitarsus covered with orange hair | karlba |
Males
1 | Forewing: hairs present in 1st medial cell and most other cells; metasomal terga T1–5 of most species with apical hair bands | 2 |
– | Forewing: hairs absent in 1st medial cell, in other cells hairs absent or restricted to radial, marginal, 1st and 2nd submarginal cells; metasomal terga T1–5 without hair bands | subgenus Asaropoda |
2 | S6 gently convex (Fig. |
3 |
– | S6 with broad depressions either side of midline (Fig. |
4 |
3 | Metasoma uniformly covered with green/bronze hair | aeruginosa |
– | Metasoma with pale blue bands, often with orange tints | chlorocyanea |
4 | Outer surface of hind tibia covered with white hair | 5 |
– | Outer surface of hind tibia covered with orange hair | 11 |
5 | Clypeus and labrum ivory or pale yellow | 6 |
– | Clypeus and labrum bright yellow | 7 |
6 | Paraocular areas without dark hairs; T2–4 hair bands broad, extending anterolaterally below the gradulus (Fig. |
murrayensis |
– | Paraocular areas with dark hairs; T2–4 apical hair bands not extending anterolaterally below the gradulus (Fig. |
pulchra |
7 | Posterior margin of S5 with distinct patch of dark, branched hair | 8 |
– | Posterior margin of S5 with at most an indistinct patch of branched hair | 9 |
8 | Lateral black marks on clypeus narrow, more than twice as long as wide; metasomal bands blue, narrow (band on T2 0.30× width of disc) | asserta |
– | Lateral black marks on clypeus about twice as long as wide; metasomal bands blue, usually with yellow tinge, band on T2 about 0.35× width of disc | adelaidae |
9 | Thoracic hair appears grey due to mixed black and pale hairs; metasomal hair bands metallic blue | walkeri |
– | Thoracic hair appears orange or pale brown; metasomal hair bands various | 10 |
10 | Metasomal bands metallic blue; S7 windows large (Fig. |
thorogoodi |
– | Metasomal bands orange or green/orange; S7 windows small, apical projection truncated (Fig. |
indistincta |
11 | Metasomal terga with dense bands of orange, scale-like hair on apical margins and similar hair distributed openly on the rest of each tergum | alpha |
– | Metasomal terga T1–T4 with coloured hair only in apical bands | 12 |
12 | Metasomal hair bands electric blue | cingulata |
– | Metasomal hair bands green or orange or a combination of these colours | 13 |
13 | Entire outer surface of hind basitarsus covered with orange hair, metasomal hair bands orange, T6 with broad orange hair band; S7 windows absent (Fig. |
paeninsulae |
– | Less than 30% of outer surface of hind basitarsus with pale hair | 14 |
14 | Thoracic hair usually bright ferruginous; S7 rounded apically (Fig. |
viridicingulata |
– | Thoracic hair pale orange; S7 with small apical projection (Fig. |
karlba |
Asaropoda Cockerell, 1926, p. 216. Type species: Saropoda bombiformis Smith, 1854 (original designation).
Length 13–24 mm; pubescence brown to grey (mostly black in aurata from New Guinea and the Bismark Archipelago); maxillary palpus with last segment fused to fifth segment so that maxillary palpus appears five-segmented.; apical margins of male metasomal sterna modified; S4 usually produced medially, rounded and having a thick brush of hair; S5 broadly and deeply emarginate; S6 shallowly emarginate medially with one or two patches of hair laterally; S7 somewhat quadrate, medioapically emarginate; S8 apically narrowed; apex of gonocoxite of male bilobed with long narrow upper lobe and small ventral lobe; gonostylus of male well developed. Forewing conspicuously hairy near costal margin, 1st discoidal cell without hairs, other closed cells with hairs sparse or absent; gonostylus of male distinct, slender, and directed apically.
Widely distributed across Australia, New Guinea and the Bismark Archipelago, but not recorded from Tasmania.
Notomegilla Brooks, 1988, p. 511.Type species: Anthophora aeruginosa Smith, 1854 (original designation).
Length 9–12 mm; metallic green/blue hair present on femur and tibia of fore and mid legs; pale paraocular marks present in males, absent in females; maxiliary palpus with 4 or 5 segments. Most or all closed cells of forewing with some rather long hairs; S6 of male simple, convex without lateral depression, edge entire to shallowly emarginate medially; apex of S7 of male greatly expanded laterally, without subapical circular area; apex of S8 narrowed, truncate or rounded.
For a full description of the subgenus see
Female. Structure. Head: wider than long; inner orbits of eyes diverging above; length f3–9 subequal. Coloration. Integument black, except ivory or pale yellow marks on labrum, mandibles, clypeus and supraclypeal area (
Male. Structure. Head: wider than long; inner orbits diverging above; length f3–10 subequal. Wings: length cu-v of hind wing subequal to second abscissa M+Cu. Metasoma: Apicomedial margin T7 bilobed. Coloration. Integument black, pale yellow marks on labrum, mandibles, clypeus, scape, paraocular and supraclypeal areas. Pubescence. Head: labrum white, gena mostly white. Legs: Forefemur posteriorly with long, light coloured hair; forecoxa white; mid and hind legs dark or black, with lighter coloured hairs on apex of femur, outer surface of tibia and basitarsus. Metasoma: T1–T6 with apical hair bands or entirely covered with adpressed hair; S6 dark. Sculpture. Head: clypeus dull with small, shallow, open punctures; interspaces pit-reticulate; Labrum somewhat shiny, with small, shallow punctures; interspaces interspaces pit-reticulate. Thorax: scutum with medium, strong, dense punctures; interspaces smooth. Metasoma: T1–T5 with shallow, close, somewhat scrobiculate punctures; interspaces pit-reticulate.
Australia.
aeruginosa and chlorocyanea.
Anthophora aeruginosa Smith, 1854, p. 336.
Amegilla aeruginosa (Smith) Michener, 1965, p. 216.
Amegilla (Notomegilla) aeruginosa (Smith) Brooks, 1988, p. 512.
Anthophora kershawi Rayment, 1944, p. 21, n. syn.
Anthophora sybilae Rayment, 1944, p. 22, n. syn.
Amegilla sybilae (Rayment) Michener, 1965, p. 217.
Amegilla (Notomegilla) sybilae (Rayment) Brooks, 1988, p. 512.
364 females and 286 males.
Lectotype of aeruginosa: male, BMNH 17B.659. The original description was from one or more male and female syntypes, reportedly collected from Hunter River, Australia. The only specimen found that was considered to be one of the syntypes was a single male bearing no collection information, but two handwritten labels: “aeruginosa Type Sm.” and “Anthophora aeruginosa Type Sm.”. Accordingly this specimen is here designated as the lectotype.
Holotype (by monotypy) of kershawi: male, Claudie River, Queensland,
Syntypes of sybilae: male, Macintosh Holding, Queensland, 14 Mar.1940,”cotype”,
Based on examination of the holotype and of Rayment’s syntypes, combined with results of DNA analyses of specimens from across its geographical range we conclude that there is evidence for only one species. Uncorrected sequence divergence was found to be 0–2.0%, which is within the limits expected for conspecific individuals (
Both sexes are easily distinguished from other Australian Amegilla by the green/bronze metallic pubescence which covers most of the dorsal surface.
Female: Pine Creek, NT, 10 Jul. 1997, Leg. R. Leijs
Length 10 mm; forewing length 9 mm.
Structure. Head: clypeus protuberant, in profile 0.4× width of eye; galea in repose reaching more than half-way between coxa of fore and mid legs; length f1 3× length f2; IOD 1.4× OOD; OS 0.6× OOD. Coloration. Yellow marks on labrum, mandibles, supraclypeal areas and inverted T-shape on clypeus; paraocular areas and scape black. Pubescence. Head: labrum and clypeus white, a light yellow hair patch in centre of paraocular area, some black hairs between antennae, near ocelli and vertex, white hair with blue-green metallic iridescence in remaining areas; gena white with metallic blue-green hair. Thorax: scutum with mixed dark and yellowish hair with a metallic blue-green iridescence; pleura pale yellow below wing base, white with metallic blue-green iridescence in other areas; thoracic sterna white with metallic blue-green iridescence; propodeum laterally, dark intermixed longer whitish hair. Legs: fore outer femur and tibia posteriorly with white hair; outer tibia and basitarsus with white hair with metallic blue-green iridescence, rest of tarsus dark; mid legs predominantly dark, white on apex of femur, white with metallic blue-green iridescence on outer surface of tibia and as a basal patch on basitarsus; hind legs predominantly dark, except white on apex of femur and white with metallic blue-green iridescence on outer surface of tibia plus a small basal patch on basitarsus; basitibial streak 0.5× length of femur. Metasoma: T2–T5: green or bronze with a metallic iridescence; T5 laterally with white hair (Fig.
Male: Pine Creek, NT, 10 July 1997, Leg. R. Leijs
Length 10 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.8× length of eye; clypeus protuberant, in profile 0.6× width of eye; galea in repose reaching more than halfway between fore and mid coxae; length f1 1.4× length f2, 0.7× length scape (excluding basal bulb) and 0.7× length f10; length f3–10 1.4× width; IOD 1.8× OOD; OS 0.9× OOD. Wings: length of marginal cell 0.8× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2.5× length second abscissa of M+Cu; length of jugal lobe about 0.6× length of vannal lobe. Metasoma: apicomedial emargination of S5 weak; S7 windows absent, median hair brush very wide at apex and narrowing towards anterior end (Fig.
The colour of the metasomal hair is predominantly green for approximately half the specimens and bronze for the remaining half, although intermediate coloration is found in a few individuals.
Month: | Jan | Feb | Mar | Apr | May | Jun | Jul | Aug | Sep | Oct | Nov | Dec |
---|---|---|---|---|---|---|---|---|---|---|---|---|
No. of records: | 16 | 19 | 31 | 37 | 67 | 55 | 25 | 39 | 58 | 60 | 126 | 70 |
Amegilla aeruginosa was observed throughout the year in the Northern parts of Australia. There were an increase in numbers at the start of the wet season, and a decrease in January and February. This may be due to a slowing down or lack of reproduction in the wettest part of the year. The strong male bias in the first two months of the year (Fig.
Widely distributed in the tropics and subtropics, with little overlap with the distribution of A. (Notomegilla) chlorocyanea (Fig.
Anthophora chlorocyanea Cockerell, 1914, p. 469.
Amegilla chlorocyanea (Cockerell) Michener, 1965, p. 216.
Amegilla (Notomegilla) chlorocyanea (Cockerell) Brooks, 1988, p. 512.
Anthophora australis Rayment, 1944, p. 24, n. syn.
Amegilla australis (Rayment) Michener, 1965, p. 216.
Amegilla (Zonamegilla) australis (Rayment) Brooks, 1988, p. 511.
Anthophora adamsella Rayment, 1944, p. 23, n. syn.
Amegilla adamsella (Rayment) Michener, 1965, p. 216.
Amegilla (Notomegilla) adamsella (Rayment, 1944) Brooks, 1988, p. 512.
Anthophora ferrisi Rayment, 1947, p. 73, n. syn.
Amegilla ferrisi (Rayment) Michener, 1965, p. 216.
Amegilla (Zonamegilla) ferrisi (Rayment) Brooks, 1988, p. 511.
Anthophora grayella Rayment, 1944, p. 27, n. syn.
Amegilla grayella (Rayment) Michener, 1965, p. 216.
Amegilla (Notomegilla) grayella (Rayment) Brooks, 1988, p. 512.
Anthophora tinsleyella jamesi Rayment, 1944, p. 30, n. syn.
Amegilla jamesi (Rayment) Michener, 1965, p. 216.
Amegilla (Notomegilla) jamesi (Rayment) Brooks, 1988, p. 512.
Anthophora luteola Rayment 1944, p. 27.
Amegilla luteola (Rayment) Michener, 1965, p. 217.
Anthophora mewiella Rayment, 1944, p. 28, n. syn.
Amegilla mewiella (Rayment) Michener, 1965, p. 217.
Amegilla (Notomegilla) mewiella (Rayment) Brooks, 1988, p. 512.
Anthophora luteola murrayi Rayment, 1944, p. 28, n. syn.
Amegilla murrayi (Rayment) Michener, 1965, p. 217.
Amegilla (Notomegilla) murrayi (Rayment) Brooks, 1988, p. 512.
Anthophora tinsleyella Rayment 1944, p. 29.
Amegilla tinsleyella (Rayment) Michener, 1965, p. 217.
1110 females and 946 males.
Holotype of chlorocyanea: female, whereabouts unknown. As the original description is unambiguous, no neotype is required.
Syntype of australis: female, Sandringham, Victoria, 7 Nov. 1936, on Dianella revoluta,
Syntype of adamsella: male, Edungalba, Queensland, May 1940, 5EEA, “Allotype”,
Syntypes of grayella: male, female, Orroroo, South Australia, 20 Feb. 1940 No. 88,
Syntypes of tinsleyellajamesi: male, female, Orroroo, South Australia, 20 Feb. 1940, 92,
Syntypes of luteola: male, female, Orroroo, South Australia, 3 Mar. 1939, “Gray 23 Adelaide”,
Syntypes of mewiella: male, Broken Hill, 20 Feb. 1940,
Syntype (presumed) of luteolamurrayi: female, Robertson, New South Wales, Feb 1940,
Syntypes (presumed) of tinsleyella: 5 males, Orroroo, SA, 4 Feb 1940, 20 Feb 1940 (3), 26 Feb 1940,
No type material was found for ferrisi but it is placed in chlorocyanea on the basis of the description provided by
The results of DNA analyses of specimens from across the complete geographical range indicated that there is only one banded Notomegilla species. Uncorrected sequence divergence was found to be 0–3.05%, which is below the usual limits for conspecific individuals. Moreover, there was no geographical pattern in the sequence variation. No morphological differences, other than colour variation, were found when Rayment’s syntypes were examined. Variations in the genitalia as illustrated by
We agree with
This species superficially resembles several Zonamegilla species, but can be distinguished by the blue/green iridescent pubescence on the fore and mid legs; females by the completely black paraocular areas and a large, dense, medial spot of pale pubescence on T5 (Fig.
Female: Sunnyside, 11km NW of Murray Bridge, 35.0536S 139.3620E, SA, 28 Dec 2003, R.Leijs & K.Hogendoorn,
Length 13 mm; forewing length 9.5 mm.
Structure. Head: clypeus protuberant, in profile 0.5× width of eye; galea in repose reaching just past forecoxa; length f1 3.5× length f2, 0.9× length scape (excluding basal bulb) and 2× length f10; f3–9 as long as wide; IOD 1.6× OOD; OS 0.8× OOD. Coloration. Pale yellow marks on labrum, mandibles, supraclypeal areas and inverted T-shape on clypeus; paraocular areas and scape black. Pubescence. Head: labrum white, remaining areas predominantly pale yellow with some black hairs between antennae and on vertex; gena white/pale brown. Thorax: scutum ginger intermixed with black hairs; pleura light ginger under wing base, remainder white; thoracic sterna white; propodeum laterally light ginger intermixed with black. Legs: foreleg brown, except white long hair posteriorly on femur and pale hair with orange or light blue metallic iridescence on outer tibia and basitarsus; mid legs black, except pale hair with orange or light blue metallic iridescence on apex of femur, outer tibia and basal part of basitarsus; hind legs black except pale with orange or light blue metallic iridescence on apex of femur, outer tibia and basal part of basitarsus; length of basitibial streak 0.4× length of femur. Metasoma: apical hair bands on T1–T4 white with metallic blue-green iridescence, margin of T4 medially with hairless shiny triangle, parts not covered by hair bands black, with some white hair medially on T4; T5 laterally with long white hair intermixed with hairs with metallic blue-green iridescence (Fig.
Male: Sunnyside, 11km NW of Murray Bridge, 35.0536S 139.36199E, SA, 28 Dec 2003, R.Leijs & K.Hogendoorn,
Length 11 mm; forewing length 9 mm.
Structure. Head: shortest distance between eyes 0.6× length of eye; clypeus protuberant, in profile 0.4× width of eye; galea in repose almost reaching mid coxa; length f1 2.4× length f2, 0.7× length scape (excluding basal bulb) and 1.1× length f11; length f3–10 1.2× width; IOD 1.8× OOD; OS 0.9× OOD. Wings: length of marginal cell 0.7× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2.3× length second abscissa of M+Cu; length of jugal lobe about 0.4× length of vannal lobe. Metasoma: apicomedial emargination of S5 weak and broad; S7 windows absent or very small (Fig.
About 10% of specimens have enough orange pigmentation to make the tergal bands and scutal hair orange and the hair of the legs pale orange. A larger number of specimens have almost white tergal bands, presumably as a consequence of wear.
Month: | Jan | Feb | Mar | Apr | May | Jun | Jul | Aug | Sep | Oct | Nov | Dec |
---|---|---|---|---|---|---|---|---|---|---|---|---|
No of records Nof 30°S: | 30 | 18 | 41 | 51 | 77 | 6 | 3 | 26 | 74 | 90 | 72 | 57 |
No of records S of 30°S: | 338 | 165 | 133 | 70 | 24 | 6 | 0 | 2 | 25 | 125 | 205 | 202 |
Amegilla chlorocyanea occurs throughout Australia, but the epicentre of the distribution is in the south of the continent, as is demonstrated by the fact that 73% of the specimens with known localities (n = 2043) have been collected south of a latitude of 30°S. The phenology changes with the latitude: peak activity is in January in the south, and in May and October in the north. In Fig.
Wide-spread throughout the arid and temperate areas of the southern part of mainland Australia and Tasmania (Fig.
Zonamegilla Popov, 1950, p. 260.
Apis zonata Linnaeus, 1758 (original designation).
Length 10–14 mm; most species with blue, green, white, or occasionally orange, metallic hair bands on metasomal terga; pale paraocular markings present in both sexes; maxiliary palpus with 6 segments; S5 of male apicomedially broadly to narrowly emarginate; S6 of male with lateral depressions on apical third, sometimes with a median protuberance, tuft of black hair apicomedially.
The following description refers to the Australian species of Zonamegilla (see
Female. Structure. Head: wider than long; inner orbits diverging above; f3–9 about equal in length. Coloration. Integument black, except yellow or pale yellow marks on labrum, mandibles, clypeus, scape, paraocular and supraclypeal area (marks are ivory in pulchra). Pubescence. Head: gena with white hairs. Legs: forecoxa and femur posteriorly light ginger in paeninsulae, white in all other species, hair of anterior face of femur and tibia white or ginger; mid and hind legs dark or black, but lighter coloured on apex of femur, on outer surface of tibia and on base of basitarsus. Metasoma: apical hair bands on T1–T4; parts not covered by hair bands dark brown or black in A. alpha, black in all other species. Punctation. Head: clypeus somewhat shiny, close to dense punctation, interspaces pit-reticulate; labrum interspaces almost smooth in murrayensis and adelaidae, reticulate in other species. Thorax: scutum with small, shallow punctures; interspaces almost smooth. Metasoma: T1–T5 somewhat shiny, with fine, shallow punctures, interspaces pit-reticulate.
Male. Structure. Head: wider than long; inner orbits diverging above; f3–10 about equal in length. Metasoma: apicomedial margin of T7 bilobed. Coloration. Integument black, except yellow or pale yellow marks on labrum, mandibles, clypeus, scape, paraocular and supraclypeal area (marks are ivory in pulchra). Pubescence. Head: gena with white hair. Thorax: sterna pale orange in paeninsulae, white in all other species. Legs: forefemur posteriorly with long, light coloured hairs; coxa greyish white in paeninsulae, white in all other species; mid and hind legs: dark or black, with lighter coloured hairs on apex of femur, outer surface of tibia and basitarsus. Metasoma: apical hair bands on T1–T5; parts not covered by hair bands dark brown in walkeri, black in all other species. S6 dark except in paeninsulae, thorogoodi and cingulata. Punctation. Head: clypeus dull, with open shallow punctures, interspaces rough pit-reticulate in karlba, pit-reticulate in all other species; labrum somewhat shiny, interspaces pit-reticulate; scutum: interspaces almost smooth. Metasoma: interspaces pit-reticulate.
India, South East Asia including southern China, and Australia.
adelaidae, alpha, asserta, cingulata, thorogoodi, indistincta, karlba, murrayensis, paeninsulae, pulchra, viridicingulata and walkeri.
Anthophora adelaidae Cockerell, 1905, p. 397.
Amegilla adelaidae (Cockerell) Michener, 1965, p. 216.
Amegilla (Zonamegilla) adelaidae (Cockerell) Brooks, 1988, p. 511.
54 females and 33 males.
Holotype of adelaidae: male, Adelaide River, NT, BMNH 17B.664.
The identity of the species adelaidae was determined unequivocally from the shape of S7 of the holotype.
Amegilla adelaidae may be recognized by the matt, pale orange tergal hair bands in both sexes; females by the broad, entire patch of white hairs on T5 (Fig.
Female: Berrimah, Research Farm Orchard, NT, 12.4333S 130.9167E, 14 May 2003, G.R. Brown & H. Wallace, DNA voucher RB266 (RL501),
Length 13 mm; forewing length 8 mm.
Structure. Head: clypeus protuberant, in profile 0.5× width of eye; galea in repose reaching beyond mid coxa; length of f1 2.7× length of f2, 0.8× length of scape (excluding basal bulb) and 1.8× length of f10; length of f3–9 0.9× width; IOD 1.3× OOD; OS 0.6× OOD. Coloration. Yellow marks on labrum, mandible, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus; f2 and apex of f1 orange ventrally. Pubescence. Head: labrum white, remaining areas predominantly pale, darker towards the vertex; black robust hairs scattered between antennae, near ocelli and on vertex, a few on clypeus; gena white, light brown towards vertex. Thorax: scutum ginger intermixed with black hair; pleura ginger with scattered black hairs under wing base, white ventrally; thoracic sterna white; propodeum laterally light ginger with scattered black hairs. Legs: forefemur posteriorly with long white hair, outer surface of foretibia and -tarsus pale yellow, inner surface of foretarsus dark; mid legs dark, except white hairs on apex of femur and on outer surface of tibia and basitarsus and a streak of contiguous short white hairs on posterior proximal part of femur; hind legs dark, except white hairs on apex of femur and outer surface of tibia, white patch on basal part of basitarsus; basitibial streak black, 0.7× length of femur. Metasoma: apical hair bands on T1–T4 white with weak light blue and orange iridescence; T5 laterally with long white hairs and few dispersed short hairs (Fig.
Male: Berrimah, Research Farm Orchard, NT, 130.9167E 12.4333S, 14 May 2003, G.R. Brown & H. Wallace, DNA voucher RB266 (RL502),
Length 12 mm; forewing length 8.5 mm.
Structure. Head: shortest distance between eyes 0.8× length of eye; clypeus protuberant, in profile 0.5× width of eye; galea in repose reaching beyond mid coxa; length of f1 1.9× length of f2, 0.6× length of scape (excluding basal bulb) and 1.1× length of f11; length of f3–10 1.1× width; IOD 1.4× OOD; OS 0.7× OOD. Wings: length of marginal cell 0.8× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2.3× length of second abscissa of M+Cu; length of jugal lobe about 0.5× length of vannal lobe. Metasoma: apicomedial emargination of S5 wide; S7 with rounded apical margin, sharp apical projection, large windows, width medial ridge 1.6× length, narrow Y-shaped brush (Fig.
Most female specimens have pale orange bands with no metallic reflections, but the bands of males are frequently paler and more often show hints of green iridescence. A few specimens, male and female, had ivory, rather than yellow face marks.
Australia, mainly in tropical and subtropical areas, including the arid zone (Fig.
Sarapoda bombiformis var α Smith, 1854, p. 318.
Saropoda alpha Cockerell, 1904, p. 204.
Amegilla (Asaropoda) alpha (Cockerell) Michener, 1965, p. 217.
5 females and 1 male.
Holotype of alpha: male, (no locality data), BMNH 17B.669.
Amegilla alpha is easily distinguished from all other Australian Zonamegilla species by the orange hair covering the dorsal surface of the metasoma in both sexes, superficially resembling that in Asaropoda species. The yellow paraocular marks and facial profile of females, the unmodified apical margins of S4 and S5 of males, and for both sexes the presence of hairs in 1st medial cell and most other cells of the forewing, separate it from Asaropoda.
Female: Jasper Gorge 54km NW Victoria River Downs, 16.02S 130.41E, NT, 30 Apr. 1974, T.Weir & T. Angeles,
Length 14 mm; forewing length 9 mm.
Structure. Head: clypeus protuberant, in profile 0.5× width of eye; galea in repose reaching just reaching mid coxa; length of f1 3× length of f2, 0.8× length of scape (excluding basal bulb) and 1.7× length of f10; f3–9 as long as wide; IOD 1.3× OOD; OS 0.5× OOD; length of marginal cell 0.8× distance from apex of marginal cell to wing tip; cu-v of hind wing approximately half length of second abscissa of M+Cu; length of vein M of hind wing 1.7 times as long as second abscissa of M+Cu; length of jugal lobe about 0.5× length of vannal lobe. Coloration. Yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus; distal part of flagellum brown ventrally from apex of f1. Pubescence. Head: labrum and clypeus with orange setae, pale yellow hairs intermixed with dark hairs on paraocular areas, frons, near ocelli and on vertex, darker near ocelli; gena white, pale ginger towards vertex. Thorax: scutum orange intermixed with black hairs; pleura orange with few black hairs under wing base, white ventrally; thoracic sterna orange; propodeum laterally orange with few black hairs. Legs: forefemur posteriorly with long white hair, outer surface of foretibia and -tarsus light orange, inner surface of tarsus brown; mid leg black, except light orange hair on apex of femur and on outer surface of tibia and basitarsus, hair on basitarsus lighter than on tibia; posterior proximal part of femur with a narrow line of light orange hair; hind legs black, except orange hair on apex of femur, scopa and basal part of basitarsus; basitibial streak brown, very short. Metasoma: apical hair bands on T1–T4 ginger with weak orange iridescence, parts not covered by hair bands dark orange; T5 laterally pale yellow (Fig.
Male: Holotype.
Length 11 mm; forewing length 7.5 mm.
Structure. Head: clypeus protuberant, in profile 0.4× width of eye; length f1 2.1× length f2, 0.6× length scape (excluding basal bulb); length f3–10 1.1× width; IOD 1.4× OOD; OS 0.7× OOD. Metasoma: apicomedial emargination of S5 wide, 40% of the sternal width, S6 with lateral depressions. Pubescence. Head: labrum white, remaining areas predominantly pale, darker towards vertex; scattered black robust hairs on clypeus laterally and in pale paraocular areas; gena white. Thorax: scutum orange-brown intermixed with black hairs; pleura light brown, white ventrally; propodeum laterally orange-brown with scattered black hairs. Legs: forefemur posteriorly with long white hairs, outer surface of tibia, tarsus orange-brown, inner surface of tarsus dark; mid legs black, except orange-brown hair on apex of femur, outer surface of tibia and basitarsus; hind legs black, except orange-brown hair on apex of femur, outer face of tibia and base of basitarsus. Metasoma: apical hair bands on T1–T5 orange-brown, elsewhere hair simple, open, orange-brown; S3–S5 medially dark, laterally orange.
(Fig.
Anthophora asserta Cockerell, 1926, p. 224.
Amegilla asserta (Cockerell) Michener, 1965, p. 216.
Amegilla (Zonamegilla) asserta (Cockerell) Brooks, 1988, p. 511.
Anthophora perasserta assertiella Rayment, 1947, p. 63, n. syn.
Amegilla assertiella (Rayment) Michener, 1965, p. 216.
Anthophora longmani Rayment, 1947, p. 21, n. syn.
Amegilla longmani (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) longmani (Rayment) Brooks, 1988, p. 511.
Anthophora perasserta Rayment, 1947, p. 62, n. syn.
Amegilla perasserta (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) perasserta (Rayment) Brooks, 1988, p. 511.
Anthophora perpulchra Rayment, 1947, p. 64, n. syn.
Amegilla perpulchra (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) perpulchra (Rayment) Brooks, 1988, p. 511.
Anthophora whiteleyella Rayment, 1947, p. 72, n. syn.
Amegilla whiteleyella (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) whiteleyella (Rayment) Brooks, 1988, p. 511.
253 females and 215 males.
Holotype of asserta: male, Lower Ferntree Gully, 22.1.1916, VIC, MV, T-11865.
Lectotype of assertiella: male, Cooranbong, NSW, 20 May 1939,
Syntype of longmani: male, Bribie Is., Queensland, “allotype”,
Syntypes of perasserta: male, female, Clermont, QLD, K.K. Spence, AM K.105230, K.105227; male, female, Edungalba, Qld, 5 Nov. 1940,
Syntypes of perpulchra: male, female, Mittagong, NSW, 2 Feb. 1940, “TYPE”and “Allotype”,
Holotype of whiteleyella (Rayment, 1947) (by presumed monotypy), male, Macquarie River, NSW, Nov. 1935,
Examination of the above type material revealed no morphological differences.
Amegilla asserta is distinguished from other Australian Zonamegilla species by a combination of the following characters: Face marks yellow; tergal hair bands pale blue. Hind tibia of females with long dark basitibial streak; T5 with dense medial patch of pale hair including a central line extending into prepygidial fimbra (Fig.
Female: Sydney Botanical gardens, 33.850S 151.200E, NSW, 31 Mar 2003, M. Bell, DNA voucher RB081 (RL490),
Length 13 mm; forewing length 9 mm.
Structure. Head: clypeus protuberant, in profile 0.4× width of eye; galea in repose reaching half-way between coxa of fore and mid legs; length of f1 3× length of f2, 0.8× length of scape (excluding basal bulb) and 1.7× length of f10; length of f3–9 0.9× width; IOD 1.3× OOD; OS 0.7× OOD. Coloration. Pale yellow-yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus. Pubescence. Head: labrum white, remaining areas predominantly pale brown with scattered black robust hairs on clypeus, paraocular areas, between antennae, near ocelli and on vertex; gena white, pale orange towards vertex. Thorax: scutum dark ginger intermixed with black hairs; pleura ginger with scattered black hairs under wing base, white ventrally; thoracic sterna white; propodeum laterally pale ginger with scattered black hairs. Legs: forefemur posteriorly with long white hair, outer surface of tibia and tarsus with white hair, inner surface of tarsus brown; mid legs dark, with whitish hair on apex of femur, posteriorly on proximal third of femur, on outer surface of tibia and forming a small basal patch posteriorly on basitarsus; hind legs black, except white hair on apex of femur and outer surface of tibia; basitibial streak black, 0.7× length of femur. Metasoma: apical hair bands on T1–T4 white with metallic blue iridescence; T5 laterally with long white hairs intermixed with short hairs, fimbria dark, medial patch as in Fig.
Male: S. of Coen, 14.0424S 143.19888E, Qld, R. Leijs & M. Batley, DNA voucher RB277 (RL777),
Length 11 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.7 length of eye; clypeus protuberant, in profile 0.5 width of eye; galea in repose almost reaching mid coxa; length of f1 2× length of f2, 0.7× length of scape (excluding basal bulb) and 1.1× length of f12; length of f3–10 1.3× width; IOD 1.2× OOD; OS 0.6× OOD. Wings: length of marginal cell 0.8× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2 times as long as second abscissa of M+Cu; length of jugal lobe about 0.3× length of vannal lobe. Metasoma: apicomedial emargination of S5 narrow; S7 windows medium size, half circular; S7 median hair brush 3× as long as wide; S7 lateral wings of median hair brush well developed an angle of ≥90° between them (Fig.
The black areas on the clypeus of males are consistently narrow, usually 4 or 5 times as long as wide, but occasionally the length is only 2.5 times the width. The colour of the tergal bands in both sexes usually displays distinct blue iridescence and appears to be less susceptible to the effects of aging than in species like pulchra. The scutal hair of both sexes is usually distinctly ginger in appearance. The colour of the flagellum varies in both sexes from dull orange-brown to dark brown on the ventral surface and from dark brown to black on the dorsal surface. Very rarely, the length of the dark streak in the hair of the hind tibia of females is 0.5× the length of the tibia.
Amegilla asserta superficially resembles thorogoodi and indistincta, but females may be distinguished by the longer hind basitibial streak the hair pattern on T5 (Fig.
Month: | Jan | Feb | Mar | Apr | May | Jun | Jul | Aug | Sep | Oct | Nov | Dec |
---|---|---|---|---|---|---|---|---|---|---|---|---|
No of records Nof 30°S: | 36 | 30 | 20 | 18 | 20 | 12 | 8 | 2 | 5 | 11 | 39 | 23 |
No of records S of 30°S: | 55 | 31 | 37 | 11 | 3 | 0 | 0 | 0 | 0 | 0 | 12 | 22 |
Amegilla asserta is one of the most common and widespread species along the eastern seaboard, reaching from the tip of Cape York into South Australia. In the south of the continent, the species is active between November and April, with a peak in January. In the north, A. asserta can be found year round, albeit at lower frequencies in July, August and September.
From Eyre Peninsula and the Lofty Ranges in South Australia, Tasmania, and the temperate areas of Victoria and New South Wales to subtropical and tropical areas along the east coast of Queensland (Fig.
Andrena cingulata Fabricius, 1775, p. 378.
Amegilla cingulata (Fabricius) Michener, 1965, p. 216.
Amegilla (Zonamegilla) cingulata (Fabricius) Brooks, 1988, p. 511.
Anthophora emendata Smith, 1879, p. 123. n. syn.
Amegilla emendata (Smith) Michener, 1965, p. 216.
Anthophora emendata gilberti Cockerell, 1905, p. 396. n. syn.
Amegilla gilberti (Cockerell) Michener, 1965, p. 216.
Anthophora lilacine Cockerell, 1921, p. 84. n. syn.
Amegilla lilacine (Cockerell) Michener, 1965, p. 216.
Anthophora fabriciana Rayment, 1947, p. 53 n. syn.
Amegilla fabriciana (Rayment) Michener, 1965, p. 216.
Amegilla (Zonamegilla) fabriciana (Rayment) Brooks, 1988, p. 511.
186 females and 148 males.
Holotype of cingulata, female, BMNH, ‘Australia’, Banks Collection, E-668712. Mr D. Notton informed us that it is stored in the Banks Collection over the cabinet label ‘Andrena cingulata Fabr. Sp. Ins. No. 17’.
Holotype of emendata (by monotypy), male, BMNH, 17B.448.
Syntype of emendatagilberti, female, QLD, BMNH, 17B.665.
Holotype of lilacine, male, Kuranda, QLD,
Holotype of fabriciana (by monotypy), female, “ No 31; In a tunnelled cell in plaster in the walls of an old house”, “Anthophoracincta, Dours”,
Based on examination of the type material, we concur with Meade-Waldo’s (1914) decision to synonimise emendata and gilberti and
Amegilla cingulata is a distinctive species with metallic blue tergal hair bands and orange scutal pubescence in both sexes. Females lack a dark basitibial dark streak on the hind legs and the disc of T5 is without pale hair (Fig.
Female: Levers Plateau, 28.33S 152.88E, Qld, 13 Mar 1966, T. F. Houston,
Length 14 mm; forewing length 9 mm.
Structure. Head: clypeus protuberant, in profile 0.3× width of eye; galea in repose reaching mid coxa; length of f1 2.9× length of f2, 0.8× length of scape (excluding basal bulb) and 1.5× length of f10; f3–9 as long as wide; IOD 1.1× OOD; OS 0.5× OOD. Coloration. Yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus; distal part of flagellum brown ventrally from f2. Pubescence. Head: labrum white, remaining areas white to light ginger, darker towards vertex and intermixed with black hairs on clypeus, paraocular areas, frons, near ocelli and on vertex; gena white, light ginger towards vertex. Thorax: scutum orange intermixed with black hairs; pleura orange with few black hairs under wing base, white ventrally; thoracic sterna white; propodeum laterally ginger with few black hairs. Legs: forefemur posteriorly with long white hairs, outer surface of fore tibia and -tarsus light ginger, inner surface of tarsus brown; mid legs black, except light ginger hairs on apex of femur and on outer surface of tibia and basitarsus, hairs on basitarsus lighter than on tibia; posterior proximal part of femur with a narrow line of light ginger hairs; hind legs black, except ginger hairs on apex of femur, scopa and basal part of basitarsus, a white tuft on apex of tibia; basitibial streak absent. Metasoma: apical hair bands on T1–T4 white with electric blue iridescence; T5 laterally white (Fig.
Male: Bombana National Park, 27.47S 153.02E, Qld, 16 Mar 1966, T. F. Houston,
Length 13 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.8 length of eye; clypeus protuberant, in profile 0.5 width of eye; galea in repose reaching mid coxa; length of f1 1.3× length of f2, 0.5× length of scape (excluding basal bulb) and 0.8× length of f11; length of f3–10 1.2× width; IOD 1.2× OOD; OS 0.5× OOD. Wings: length of marginal cell 0.9× distance from apex of marginal cell to wing tip; vein M of hind wing 2.3 times as long as second abscissa of M+Cu; length of jugal lobe about 0.5× length of vannal lobe. Metasoma: apicomedial emargination of S5 intermediate width and depth; S7 windows medium size, median hair brush 4× width, lateral wings of hair brush narrow almost perpendicular to the long axis (Fig.
The thoracic hair, especially of females, is consistently bright orange, but the metasomal bands are seldom infused with orange and even then, the colour is mostly restricted to the band on T1. The colour of the flagellum is variable, as in asserta. Two females were found with a few white hairs on T5 forming the narrowest of longitudinal lines.
Along the east coast of New South Wales and Queensland (Fig.
40 females and 27 males.
Holotype: female: Millstream Falls, 17.6427S 145.4588E, 4 Jul 2007, R. Leijs & M. Batley, DNA voucher RB312 (RL867),
Allotype: male, Iron Range, QLD, 1 Jul 2007, 12.7465S 143.2556E, R.Leijs & M. Batley,
Paratypes: male, female, 23 km SW of Agnes Water, QLD, 24.3500S 151.9333E, 31 Jan 2007, M. Batley, DNA vouchers RB194, RB195, AM K-290886, AM K-290887.
Amegilla indistincta is distinguished from other Australian Zonamegilla species by the following combination of characters: Tergal hair bands usually with a yellowish tint. Hind tibia of females with a short dark streak (≤ 0.4× length hind tibia); pale hair on T5 forming a relatively small medial patch (Fig.
Female: holotype.
Length 14 mm; forewing length 9 mm.
Structure. Head: clypeus protuberant, in profile 0.4× width of eye; galea in repose just reaching mid coxa; length of f1 3× length of f2, 0.9× length of scape (excluding basal bulb) and 1.6× length of f10; length of f3–9 0.9× width; IOD 1.3× OOD; OS 0.6× OOD; length of marginal cell 0.8× distance from apex of marginal cell to wing tip; length of cu-v of hind wing approximately half the length of second abscissa of M+Cu; length of vein M of hind wing 2.2× length of second abscissa of M+Cu; length of jugal lobe about 0.5× length of vannal lobe. Coloration. Yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus; small brown spot on f2. Pubescence. Head: labrum and clypeus white, paraocular areas and frons white intermixed with black hairs, light ginger intermixed with black hairs near ocelli and on vertex; gena white, light ginger towards vertex. Thorax: scutum light orange intermixed with black hair; pleura light orange with few black hairs under wing base, white ventrally; thoracic sterna white; propodeum laterally light orange with few black hairs. Legs: forefemur posteriorly with long white hair, outer surface of tibia and tarsus greyish white, inner surface of tarsus brown; mid legs dark, except white hair on apex of femur and on outer surface of tibia and basal part of basitarsus; apex of tibia with brown spot; posterior proximal part of femur with narrow line of white hair; hind legs black, except greyish white hair on apex of femur, scopa, white hair on basal part of basitarsus and on apex of tibia; basitibial streak black, 0.4× length of femur. Metasoma: apical hair bands on T1–T4 greyish white with green-blue iridescence; T5 laterally white, fimbria dark brown, medial patch of dispersed short white hairs around a denser longitudinal line (Fig.
Male: allotype.
Length 12 mm; forewing length 9 mm.
Structure. Head: shortest distance between eyes 0.5× length of eye; clypeus protuberant, in profile 0.6× width of eye; galea in repose reaching mid coxa; length of f1 1.5× length of f2, 0.6× length of scape (excluding basal bulb) and 0.8× length of f1; f3–10 as long as wide; IOD 1.3× OOD; OS 0.6× OOD. Wings: length of marginal cell 0.8× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2.1× length of second abscissa of M+Cu; length of jugal lobe about 0.4× length of vannal lobe. Metasoma: apicomedial emargination of S5 moderately wide and shallow; S7 with wide medial ridge and truncate apical projection, hair pattern almost inverted T-shaped, apical half weak (Fig.
In the subtropics and tropics along the east coast of Queensland (Fig.
The specific epithet refers to the fact that specimens of this species were found among the type series of other species described by Rayment (indistincta in Latin means ’not distinguished’).
Amegilla indistincta is closely related to karlba and superficially resembles thorogoodi and asserta. Females may be distinguished from asserta by the length of the hind tibial streak and from karlba and thorogoodi by the relatively small patch of pale hair on T5. Males may be distinguished from asserta by the absence of a hair patchon S5, and from thorogoodi by the shape of S7 (Fig.
Males of indistincta are not easily distinguishable from those of karlba, but the small number of specimens available all had a truncate apical projection on S7 (Fig.
28 females and 19 males.
Holotype: female, 12km NNW of Mt Cahill, NT, 12.46S 132.39E, 20 Jun 1973, T. Weir & T. Angeles,
Allotype: male, 19km NE by E of Mt Cahill, NT, 12.47S 132.51E, 16.xi.1972, T. Weir & A. Allwood,
Paratypes: male, 16km E by N of Mt Cahill, NT, 12.8333S 132.8500E, 16.xi.1972, T. Weir & A. Allwood,
Amegilla karlba is distinguished from other Australian Zonamegilla species by the following characters: Metasomal hair bands of both sexes yellow ochre coloured; hair on outer face of the hind tibia usually orange or brown. Basitibial hair streak on hind leg of females short; hind basitarsus less than half covered with pale hair; T5 with pale hair reaching lateral margins. S7 of males with a broad medial ridge resulting in small, lightly pigmented windows and a broad but distinct apical projection (Fig.
Female: holotype.
Length 13 mm; forewing length 9 mm.
Structure. Head: clypeus protuberant, in profile 0.3× width of eye; galea in repose reaching mid coxa; length of f1 3.1× length of f2, 0.7× length of scape (excluding basal bulb) and 1.6× length of f10; f3–9 as long as wide; IOD 1.2× OOD; OS 0.6× OOD; length of marginal cell 0.8× distance from apex of marginal cell to wing tip; cu-v of hind wing 2.7× length of second abscissa of M+Cu; length of vein M of hind wing 2× length of second abscissa of M+Cu; length of jugal lobe about 2× length of vannal lobe. Coloration. Yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus; distal part of flagellum orange-brown ventrally from f2. Pubescence. Head: labrum white, remaining areas predominantly pale yellow, darker towards vertex; scattered black robust hairs on clypeus, paraocular areas, frons, near ocelli and on vertex; gena white, pale yellow towards vertex. Thorax: scutum ginger intermixed with black hair; pleura ginger under wing base, turning white ventrally; thoracic sterna pale brown; propodeum laterally ginger with scattered black hairs. Legs: forefemur posteriorly with long white hair, outer surface of foretibia and -tarsus light brown, inner surface of tarsus brown; mid legs dark, except pale brown hair on apex of femur and outer surface of tibia, slightly darker than on foretibia, posteriorly proximal end of femur with narrow line of white hair, apex of tibia with brown spot, basitarsus white basally; hind legs black, except pale brown hair on apex of femur and outer surface of tibia, a patch of pale brown hair on base of basitarsus, small white tuft on apex of tibia; basitibial streak brown, 0.25× length of femur. Metasoma: apical hair bands on T1–T4 yellow ochre with orange and weak light blue iridescence; T5 laterally with moderately dense white hairs, fimbria brown; T5 entirely covered with open pale yellow ochre hairs, a denser longitudinal line extends into fimbria (Fig.
Male: allotype.
Length 11 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.7× length of eye; clypeus protuberant, in profile 0.6× width of eye; galea in repose reaching mid coxa; length of f1 1.8× length of f2, 0.5× length of scape (excluding basal bulb) and 0.9× length of f11; length of f3–10 1.3× width; IOD 1.4× OOD; OS 0.7× OOD. Wings: length of marginal cell 0.8× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2.2× length of second abscissa of M+Cu; length of jugal lobe about 0.4× length of vannal lobe. Metasoma: apicomedial emargination of S5 very shallow, moderately wide; S7 with very broad medial ridge leaving small, lightly pigmented windows, apical projection very broad, but distinct and not truncate, hair pattern almost inverted T-shaped (Fig.
Arnhem Land, Kakadu, Kimberleys (Fig.
The specific epithet is a noun in apposition referring to the colour of the tergal hair bands (karlba in the language of the Kuninjku people of Western Arnhemland means yellow ochre (
Amegilla karlba is similar to indistincta and, to a lesser extent, viridicingulata. Females may be distinguished from viridicingulata by the extent of pale hair on the hind basitarsus and from indistincta the more extensive area of white hair on T5 (Fig.
Anthophora murrayensis Rayment, 1939, p. 288.
Amegilla murrayensis (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) murrayensis (Rayment) Brooks, 1988, p. 511.
Anthophora longula Rayment, 1947, p. 59. n. syn.
Amegilla longula (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) longula (Rayment) Brooks, 1988, p. 511.
Anthophora subsalteri Rayment, 1947, p. 69. n. syn.
Amegilla subsalteri (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) subsalteri (Rayment) Brooks, 1988, p. 511.
229 females and 132males.
Syntypes of murrayensis, male, female, Gunbower, VIC, 16 Mar. 1940, No. G500, “Type” & “allotype”,
Syntypes of longula, male, female, Orroroo, SA, 3 & 10 Feb. 1940,
The results of DNA analyses of specimens from across the complete geographical range showed no geographical pattern with respect to sequence variation. The uncorrected sequence divergence was found to be 0–1.3% (Table
The syntypes of murrayensis and longula were examined and considered to be conspecific. Type material for subsalteri, presumed to be the holotype by monotypy, was not found, but from Rayment’s description and drawings (
Amegilla murrayensis is a relatively small species with pale yellow face marks and narrow (about 0.3x the width of the disc on T2) apical hair bands which are usually pale blue, occasionally with an orange tint, but never bright orange; female hind tibia with a dark streak at least 0.5× as long as the tibia; T5 with a patch of scattered white hair that narrows laterally and with a longitudinal line of denser white hair that does not intrude significantly into the prepygidial fimbria. Both sexes can be distinguished from other species by hair bands on T1–4 that appear broader laterally below the gradulus because of numerous scattered pale hairs on the disc (Figs
Female: Sunnyside, N of Murray Bridge, 35.0500S 139.3600E, 27 Feb 2003, R.Leijs & K. Hogendoorn,
Length 12 mm; forewing length 8 mm.
Structure. Head: clypeus protuberant, in profile 0.5 width of eye; galea in repose reaching half-way between fore and mid coxae; length of f1 2.8× length of f2, 0.9× length of scape (excluding basal bulb) and 1.6× length of f10; length of f3–9 0.9× width; IOD 1.4× OOD; OS 0.8× OOD. Coloration. Yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus. Pubescence. Head: labrum white, remaining areas predominantly pale yellow with scattered black robust hairs on clypeus, paraocular areas, between antennae, near ocelli and on vertex; gena white. Thorax: scutum ginger intermixed with black hairs; pleura ginger with scattered black hairs under wing base, white ventrally; thoracic sterna white; propodeum laterally light ginger with scattered black hairs. Legs: forefemur posteriorly with long white hairs, outer surface of fore tibia and tarsus white, inner surface of tarsus dark; mid legs dark, except white hairs on apex of the femur, posteriorly on proximal one third of femur and on outer surface of tibia and basitarsus; hind legs black, except white hairs on apex of femur and outer surface of tibia; basitibial streak black, 0.6–0.9 length of femur. Metasoma: apical hair bands on T1–T4 white with iridescence varying from light blue to greenish orange; T5 laterally with long white hairs and few dispersed short hairs (Fig.
Male: Coen, 13.94415S 143.20022E, QLD, 27 June 2007,
Length 11 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.7 length of eye; clypeus protuberant, in profile 0.5 width of eye; galea in repose reaching halfway between fore and mid coxae; length of f1 1.5× length of f2, 0.5× length of scape (excluding basal bulb) and 0.8× length of f11; length of f3–10 1.2× width; IOD 1.5× OOD; OS 0.7× OOD. Wings: length of marginal cell 0.8× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2.6 times as long as second abscissa of M+Cu; length of jugal lobe about 0.5× length of vannal lobe. Metasoma: apicomedial emargination of S5 wide and deep; S7 windows small, median hair brush 3–4× width, lateral wings of hair brush well developed making an angle of 60° with long axis of brush (Fig.
Males from southern Western Australia often have more black hairs on the clypeus and paraclypeal areas than specimens from northern Queensland or specimens from the Lofty Ranges in South Australia. There is also some variation in the width of the pale patch on T5 in fresh females. Some specimens from the NW Pilbarra and Barrow Island are seemingly larger, have wider tergal bands and a more intense patch of pale hairs on female T5 and have almost ivory face marks and white apical bands on T1–4. These specimens are sufficiently different from murrayensis and may belong to an undescribed species. Future collections and molecular work may shed light on their identity.
There were some problems with the molecular delineation of murrayensis and pulchra, as mentioned in the Results and Discussion section. The sequences obtained with the M202/M70 primers resulted in two morphologically similar clades of murrayensis specimens, often with specimens from the same localities in different clades (Suppl. material
Month: | Jan | Feb | Mar | Apr | May | Jun | Jul | Aug | Sep | Oct | Nov | Dec |
---|---|---|---|---|---|---|---|---|---|---|---|---|
No of records N of 30°S: | 10 | 6 | 24 | 10 | 24 | 4 | 7 | 25 | 4 | 11 | 17 | 5 |
No of records S of 30°S: | 161 | 16 | 6 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 7 | 16 |
Amegilla murrayensis is the most widespread and common species in the subgenus Zonamegilla. Below 30°S the species is active from November until March, with a peak in January. In the north of the continent they can be found year round possibly with a peak in May.
Widespread throughout Australia, but not found in Tasmania (Fig.
26 females and 40 males.
Holotype: female, N of Bamaga, Qld, 10.84117S 142.42316E, 28 Jun 2007, R. Leijs & M. Batley, DNA voucher RB287 (RL795),
Allotype: male, N of Bamaga, Qld, 10.84117S 142.42316E, 28 Jun 2007, R. Leijs & M. Batley, DNA voucher RB286 (RL793),
Paratypes: 2 males, 2 females, same locality data as holotype,
Amegilla paeninsulae is a distinctive species with orange tergal hair bands, sometimes with green iridescence, orange pubescence on the scutum, orange scopa on the hind legs and no dark basitibial streak in females. Females have T5 covered with scattered orange hair and an orange medial streak (Fig.
Female: holotype
Length 14 mm; forewing length 9 mm.
Structure. Head: clypeus protuberant, in profile 0.4× width of eye; galea in repose reaching just past forecoxa; length of f1 3× length of f2, equal to length of scape (excluding basal bulb) and 1.4× length of f10; f3–9 as long as wide; IOD 1.5× OOD; OS 0.5× OOD. Coloration. Yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas, inverted T-shape on clypeus; distal part of flagellum brown ventrally from f2 onwards. Pubescence. Head: labrum white, clypeus light ginger intermixed with black, hairs in paraocular areas, frons, near ocelli and on vertex, ginger hair darker towards the top; gena white, light ginger towards vertex. Thorax: scutum ginger intermixed with black hair; pleura ginger with scattered black hair under wing base, light ginger ventrally; thoracic sterna light ginger; propodeum laterally ginger with scattered black hairs. Legs: forefemur posteriorly with long, light ginger hair intermixed with some black hairs, outer surface of foretibia and -tarsus light ginger, inner surface of tarsus brown, coxa light ginger; mid legs black, except light ginger hair on apex of femur and on outer surface of tibia and basitarsus and a small patch of light ginger hair on posterior proximal part of femur; hind legs black, except ginger on apex of femur, posterior rim of outer surface of tibia and basal part of basitarsus; basitibial streak absent. Metasoma: apical hair bands on T1–T4 orange-brown with orange and green iridescence; T5 laterally with moderately long light ginger hairs (Fig.
Male: allotype.
Length 12 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.7× length of eye; clypeus protuberant, in profile 0.5× width of eye; galea in repose reaching past mid coxa; length of f1 1.6× length of f2, 0.5× length of scape (excluding basal bulb) and 0.9× length of f11; length of f3–10 1.3× width; IOD 1.3× OOD; OS 0.6× OOD. Wings: length of marginal cell 0.9× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2.1× length second abscissa of M+Cu; length of jugal lobe about 0.4× length of vannal lobe. Metasoma: apicomedial emargination of S5 wide and deep; S7 windows absent; S7 median hair brush, broad 2× width; S7 with a small rounded head, very wide medial ridge leaving no lightly pigmented windows, no apical projection and a flattened apical margin (Fig.
Most males have predominantly orange tergal bands, while those of females are usually a mixture of iridescent green and orange. The hair on the sterna and coxae of most males is pale orange and some males lack the medial brown patch on T6.
In tropical rainforest patches on Cape York, Queensland (Fig.
The specific epithet refers to its distribution on Cape York Peninsula.
Anthophora pulchra Smith, 1854, p. 335.
Amegilla pulchra (Smith) Michener, 1965, p. 217.
Amegilla (Zonamegilla) pulchra (Smith) Brooks, 1988, p. 511.
Anthophora holmesi Rayment, 1947, p. 56, n. syn.
Amegilla holmesi (Rayment) Michener, 1965, p. 216.
Amegilla (Zonamegilla) holmesi (Rayment) Brooks, 1988, p. 511.
Anthophora parapulchra Rayment, 1947, p.61, n. syn.
Amegilla parapulchra (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) parapulchra (Rayment) Brooks, 1988, p. 511.
Anthophora salteri Cockerell, 1905, p. 398, n. syn.
Amegilla salteri (Cockerell) Michener, 1965, p. 217.
Amegilla (Zonamegilla) salteri (Cockerell) Brooks, 1988, p. 511.
Anthophora pulchra townleyella Rayment, 1947, p. 67. n. syn.
Amegilla townleyella (Rayment) Michener, 1965, p. 217.
Anthophora shafferyella Rayment, 1947, p. 70. n. syn.
Amegilla shafferyella (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) shafferyella (Rayment) Brooks, 1988, p. 511.
Anthophora perpulchra wallaciella Rayment, 1947, p. 65. n. syn.
Amegilla wallaciella (Rayment) Michener, 1965, p. 217.
171 females and 182 males.
Lectotype of pulchra, male, “pulchra type Sm.”, “Anthophora pulchra type” BMNH 17B.666b, here designated.
Another, female, specimen in the British Museum bore the following labels: Moreton Bay; pulchra type ♀ Sm; Anthophora pulchra type; Amegilla niveocincta SM D.B. Baker 2008: BMNH 17B.666a. As the specimen was unlike any Australian species, we have no reason to doubt Baker’s identification and the corollary that there has been a labelling error.
Syntype of townleyella, female, Lismore, NSW, 8 Feb 1940, “Type, Anthophora salteritownleyella”,
Lectotype of parapulchra, female, Hunters Hill, Sydney, Dec. 1939, “No 28”, “Type, Anthophora parapulchra”,
Syntypes of holmesi: female, Como, NSW, 4 Apr. 1940, “Type, Anthophora holmesi”,
Holotype of salteri (by monotypy): male, N.S.Wales, BMNH 17B.665.
Holotype of shaffereyella (by monotypy): male, Mossman, Queensland, Feb. 1940, Anthophora salterishafferyella,
Lectotype of perpulchrawallaciella: female, Hunters Hill, NSW, 20 Mar. 1940, “type, Anthophora perpulchra wallaciella”,
Examination of the above types indicated that holmesi, parapulchra, salteri and townleyella were conspecific with pulchra. The holotype of shafferyella had a dense hair patch on S5 an orange tint in the tergal hair bands like adelaidae, as suggested by
For diagnosis and description we used specimens from the Sydney area because they vary less compared to those from the Brisbane area (see also under variation and remarks)
Amegilla pulchra is a species with ivory face marks and paraocular areas with some long dark hairs. Tergites with pale blue or white hair bands that are not broadened laterally below the lateral arm of the gradulus (Figs
Female: East Kurrajong, 33.500S 150.767E, NSW, 8 Jan 2003, R Spooner Hart, DNA voucher RB083 (RL494),
Length 14 mm; forewing length 9 mm.
Structure. Head: clypeus protuberant, in profile 0.4× width of eye; galea in repose reaching half-way between coxa of fore and mid legs; length of f1 2.8× length of f2, 0.8× length of scape (excluding basal bulb) and 1.6× length of f10; length of f3–9 0.9× width; IOD 1.2× OOD; OS 0.6× OOD. Coloration. Ivory marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus. Pubescence. Head: labrum white, remaining areas predominantly pale, darker towards vertex with scattered black robust hairs on clypeus, paraocular areas, between antennae, near ocelli and on vertex; gena white, ginger towards vertex. Thorax: scutum ginger intermixed with many black hairs, therefore overall darker than other species; pleura ginger with scattered black hair under wing base, white ventrally; thoracic sterna white; propodeum laterally ginger with scattered black hair. Legs: forefemur posteriorly with long white hair, outer surface of foretibia and -tarsus greyish white, inner surface of tarsus dark; mid legs black, except white hair on apex of femur and on outer surface of tibia and basitarsus, contiguous short white hairs on posterior proximal part of femur; hind legs black except white hair on apex of femur and outer surface of tibia, very small white patch on basal part of basitarsus; basitibial streak black, 0.8× length of femur. Metasoma: apical hair bands on T1–T4 white with very weak light blue iridescence; T5 laterally with moderately long white hair (Fig.
Male: Northbridge, 33.800S 151.217E, NSW, 27 Feb 2003, M. Bell, DNA voucher RB078 (RL487),
Length 12 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.5× length of eye; clypeus protuberant, in profile 0.5× width of eye; galea in repose reaching just past forecoxa; length of f1 2× length of f2, 0.6× length of scape (excluding basal bulb) and 1.1× length of f11; length of f3–10 1.2× width; IOD 1.4× OOD; OS 0.7× OOD. Wings: length of marginal cell 0.8× distance from apex of marginal cell to wing tip; length of vein M of hind wing 2.8× length second abscissa of M+Cu; length of jugal lobe about 0.4× length of vannal lobe. Metasoma: apicomedial emargination of S5 narrow and deep; S7 windows medium size, median hair brush 3× width, lateral wings of hair brush narrow but well developed with 110° angle between them (Fig.
Most specimens of pulchra in collections have relatively narrow white bands with small amounts of green-blue iridescence and ivory face markings. However, examination of a series of fresh specimens collected from the Brisbane area on two consecutive days showed iridescent bands that varied in colour from green-blue to orange or white. Some specimens also had yellowish face marks and specimens varied with respect to the shape of the white patch on female T5, some approaching those found in murrayensis. There was however no correlation with sequenced mitochondrial DNA, because the majority of those specimens shared the same mitochondrial haplotypes (Suppl. material
In all phylogenetic analyses murrayensis and pulchra appeared as sister species (Fig.
Mainly east of the Great Dividing Range in New South Wales and Queensland (Fig.
Anthophora thorogoodi Rayment, 1939, p. 289.
Amegilla thorogoodi (Rayment) Michener, 1965, p. 217.
Amegilla (Zonamegilla) thorogoodi (Rayment) Brooks, 1988, p. 511.
95 females and 68 males.
Holotype of thorogoodi: male, Proserpine, QLD, 15 Nov. 1937,
Amegilla thorogoodi is distinguished from other Australian Zonamegilla species by the following characters: Scutal hair of both sexes brown; apical tergal hair bands predominantly blue. Hind basitibial streak of females short; pale pubescence of T5 forming a large medial patch (Fig.
Female: Iron Range, 12.743S 143.2352E, 1 Jul 2007, R. Leijs & M. Batley, DNA voucher RB302 (RL838),
Length 13.5 mm; forewing length 9 mm.
Structure. Head: shortest distance between eyes equal to the length of the eye; clypeus protuberant, in profile 0.4× width of eye; galea in repose reaching just reaching mid coxa; length of f1 3.2× length of f2, 0.8× length of scape (excluding basal bulb) and 1.5× length of f10; length of f3–9 0.8× width; IOD 1.6× OOD; OS 0.5× OOD. Coloration. Black except labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas with pale yellow marks; pale mark on clypeus inverted T-shaped. Pubescence. Head: labrum white, remaining areas pale yellow, darker towards vertex with scattered black robust hairs on clypeus, paraocular areas, between antennae, near ocelli and on vertex; gena white, ginger towards vertex. Thorax: scutum ginger intermixed with black hair; pleura ginger with scattered black hair under wing base, white ventrally; thoracic sterna white; propodeum laterally light ginger with scattered black hairs. Legs: forefemur posteriorly with long white hair, foretibia and -tarsus whitish on outer surface, dark on inner surface; mid legs black, except greyish white hair on apex of femur and on outer surface of tibia and basitarsus and a dense patch of short white hair on posterior proximal part of femur; hind legs black, except white hair on apex of femur; tibia white on posterior rim, greyish-white with orange tinge in scopal area and black anteriorly; whitish patch on basal part of basitarsus; basitibial streak black, 0.3x length of femur. Metasoma: apical hair bands on margin T1–T4 iridescent blue with orange tinge, especially across anterior edge; T5 laterally with moderately long white hairs (Fig.
Male: Bloomfield near rubbish tip, 15.9011S 143.34161E, Qld, 2 Jul 2007, DNA voucher RB311 (RL865),
Length 13 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.7× length of eye; clypeus protuberant, in profile 0.4× width of eye; galea in repose reaching halfway between fore and mid coxae; length of f1 2.1× length of f2, 0.5× length of scape (excluding basal bulb) and as long f11; length of f3–10 1.3× width; IOD 1.6× OOD; OS 0.7× OOD. Wings: length of marginal cell 0.9× distance from apex of marginal cell to wing tip; length of vein M of hind wing 1.8× length of second abscissa of M+Cu; length of jugal lobe about 0.4× length of vannal lobe. Metasoma: apicomedial emargination of S5 wide and very shallow; S7 head large, medial ridge moderately narrow resulting in a narrow apical projection, large weakly pigmented windows and an inverted Y-shaped brush (Fig.
Variation in the colour of the tergal bands caused by fading of the colour and varying amounts of orange can complicate identification, but no other significant variation was observed. Only two specimens had scutal hair with a grey appearance.
In the subtropics and tropics along the east coast of Queensland (Fig.
Closely related to the allopatric species walkeri, but easily distinguished by the brown scutal hair in both sexes. Similar to A. asserta and A. indistincta, but females distinguished by the length of the hind basitibial streak. Males may be distinguished from those of asserta by the absence of a hair patch on S5 and from indistincta by the colour of the tergal hair bands and the shape of S7 (Fig.
16 females and 27males.
Holotype: Female, Cooktown, Qld, 15.4898S 145.2413E, 3 Jul 2007, R. Leijs & M. Batley, DNA voucher RB308 (RL857),
Allotype: Cooktown, Qld, 15.4898S 145.2413E, 3 Jul 2007, R. Leijs & M. Batley, DNA voucher RB309 (RL859),
Paratypes: 4 males, 3 females, same locality data as holotype,
Amegilla viridicingulata is a distinctive species with orange-brown scutal pubescence, tergal hair bands with green iridescence and orange hair on the hind legs in both sexes. Females have a hind tibial scopa without a dark streak and T5 with a broad area of scattered white hair above the fimbria (Fig.
Female: holotype.
Length 12 mm; forewing length 8.5 mm.
Structure. Head: clypeus protuberant, in profile 0.44× width of eye; galea in repose reaching reaching mid coxa; length of f1 4× length of f2, 0.9× length of scape (excluding basal bulb) and 1.6× length of f10; f3–9 as long as wide; IOD 1.3× OOD; OS 0.5× OOD. Coloration. Pale yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas, inverted T-shape on clypeus; f2 red-brown ventrally, remainder of flagellum brown ventrally. Pubescence. Head: labrum white, light ginger intermixed with black hair on clypeus, paraocular areas, frons, near ocelli and on vertex, gena white, light ginger towards vertex. Thorax: scutum ginger intermixed with black hair; pleura ginger with few black hairs under wing base, white ventrally; thoracic sterna light ginger; propodeum laterally ginger with few black hairs. Legs: forefemur posteriorly with long white hair, outer surface of tibia and tarsus light ginger, inner surface of tarsus brown; mid legs black, except light ginger hair on apex of femur and on outer surface of tibia and basitarsus and a narrow line of light ginger hairs on posterior proximal part of femur; hind legs black, except ginger hair on apex of femur, posterior rim of the outer surface of tibia and basal part of basitarsus, a white tuft on the apex of tibia; basitibial streak absent. Metasoma: apical hair bands on T1–T4 light orange-brown with clear green iridescence; T5 laterally white, fimbria brown, medial patch well developed, centrally white, laterally greyish-white and narrowing, medial streak overlapping fimbria (Fig.
Male: allotype.
Length 11 mm; forewing length 8 mm.
Structure. Head: shortest distance between eyes 0.7× length of eye; clypeus protuberant, in profile 0.5× width of eye; galea in repose reaching halfway between fore and mid coxae; length of f1 1.6× length of f2, 0.5× length of scape (excluding basal bulb) and 0.8× length of f11; length of f3–10 1.3× width; IOD 1.3× OOD; OS 0.5× OOD. Wings: length of marginal cell equal to distance from apex of marginal cell to wing tip; length of vein M of hind wing 1.6× length second abscissa of M+Cu; length of jugal lobe about 0.3× length of vannal lobe. Metasoma: apicomedial emargination of S5 wide and shallow; S7 with very wide medial ridge, resulting in a flattened apex with no apical projection, very small weakly pigmented areas and an inverted T-shaped brush, weak and broadened towards the apex (Fig.
Some males had white metasomal hair bands with blue reflections, making them difficult to recognise before dissection.
Coastal NE Queensland (Fig.
The specific epithet is a Latin adjective meaning green banded.
Anthophora walkeri Cockerell, 1905, p. 396.
Amegilla walkeri (Cockerell) Michener, 1965, p. 217.
Amegilla (Zonamegilla) walkeri (Cockerell) Brooks, 1988, p. 511.
Anthophora darwini Cockerell, 1910, p. 409. n. syn.
Amegilla darwini (Cockerell) Michener, 1965, p. 216.
Holotype of walkeri, female, Baudin I. Long Reef, WA, 91-155, 4593, BMNH 17B.663.
Holotype of darwini, male, P. Darwin, Turner Coll. 1910-7, 11-02, “Anthophora darwini, Ckll”, BMNH 17B.448.
The holotype of darwini bears a label “Amegilla=walkeri, M.A. Lieftinck, 1958”. Examination the type specimens confirmed Lieftinck’s and Brook’s (1988) decisions to synonymise darwini with walkeri.
66 females and 63 males.
Both sexes of walkeri have grey pubescence on the scutum due of a mixture of black and white hair and conspicuous light blue, metallic hair bands on the terga. Females have a broad band of white hair bordering the fimbria on T5 (Fig.
Female: Darwin, East Point, 28 Feb.2006, 12.4130S 130.8300E, D. A. Young,
Length 12 mm; forewing length 8.5 mm.
Structure. Head: clypeus protuberant, in profile 0.4× width of eye; galea in repose reaching just past fore coxa; length of f1 2.6× length of f2, 0.7× length of scape (excluding basal bulb) and 1.6× length of f10; length of f3–9 1.1× width; IOD 1.3× OOD; OS 0.5× OOD. Coloration. Yellow marks on labrum, mandibles, scape, clypeus, paraocular and supraclypeal areas; inverted T-shape on clypeus; f2 orange, f3–10 brown ventrally. Pubescence. Head: white, intermixed with black hair on clypeus, paraocular areas, frons, near ocelli and on vertex; gena white. Thorax: scutum white intermixed with black hair, producing an overall grey appearance; pleura white with scattered black hair under wing base; thoracic sterna white; propodeum laterally white with scattered black hair. Legs: fore femur posteriorly with long white hair, outer surface of tibia and tarsus white, inner surface of tarsus dark; mid legs black, except white hair on the apex of femur and on outer surface of tibia and basitarsus and a dense streak of short white hair on posterior proximal part of femur; hind legs black, except white hair on apex of femur, posterior rim of the outer surface of tibia, white patch on basal part of basitarsus; basitibial streak black, 0.4× length of femur. Metasoma: apical hair bands on T1–T4 white with clear light blue iridescence; T5 laterally with moderately long white hair (Fig.
Male: Darwin, East Point, 28 Feb.2006, 12.4130S 130.8300E, D. A. Young,
Length 10 mm; forewing length 7 mm.
Structure. Head: shortest distance between eyes 0.7× length of eye; clypeus protuberant, in profile 0.5× width of eye; galea in repose reaching just past mid coxa; length of f1 1.8× length of f2, 0.9× length of scape (excluding basal bulb) and 0.9× length of f11; length of f3–10 1.2× width; IOD 1.3× OOD; OS 0.6× OOD. Wings: length of marginal cell 0.9× distance from apex of marginal cell to wing tip; length of vein M of hind wing 1.6× length of second abscissa of M+Cu; length of jugal lobe about 0.4× length of vannal lobe. Metasoma: apicomedial emargination of S5 wide and shallow; S7 head large, medial ridge moderately narrow resulting in a narrow, rounded apical projection, large weakly pigmented windows and an inverted Y-shaped brush (Fig.
In tropical areas of the Northern Territories and Western Australia (Kimberleys) (Fig.
Closely related to the allopatric species thorogoodi from which it can be distinguished by the colour of the thoracic hair.
R.L., M.B. and K.H. designed the research, collected and examined specimens and wrote the paper, R.L. extracted and sequenced DNA and interpreted the molecular data.
The authors thank Melissa Bell for donating a large collection of ethanol preserved specimens from around Australia collected by the following persons: M. Eden, J. Klumpp, W. Forno, G. Hambridge, R. Spooner Hart and R.Storey. The authors also acknowledge Chris Brocks, Graham Brown, Ralph Foster, Brenda Kranz, Wojciech Pulawski, H. Wallace, Geoff Williams and Andy Young for providing fresh specimens preserved in ethanol. Jacqui Street performed part of the DNA analysis and photography. This work was supported by an ABRS grant to RL, MB and KH.
Phylogenetic trees and table of DNA voucher numbers, including Genbank accession numbers
Data type: phylogenetic data
Explanation note: Figure S1.CO1 (primers M70/M202) neighbour-joining tree calculated using uncorrected sequence divergence in PAUP*. Terminals are labelled with RB-numbers, which refer to Table S1, sequencing primer or ‘cons’, indicating consensus sequence based on forward and reverse sequencing.
Figure S2.CO1 (primers M414/M423) neighbour-joining tree calculated using uncorrected sequence divergence in PAUP*. Terminals are labelled with RB-numbers, which refer to Table S1, sequencing primer or ‘cons’, indicating consensus sequence based on forward and reverse sequencing.
Table S1. Table of DNA specimens and GenBank accession numbers and locality data.
Examined specimens
Data type: specimens database
Explanation note: The database contains information copied from specimen labels for each species: including repository, catalogue numbers and DNA voucher numbers.