Research Article |
Corresponding author: Michael C. Orr ( michael.christopher.orr@gmail.com ) Corresponding author: Chao-Dong Zhu ( zhucd@ioz.ac.cn ) Academic editor: Jack Neff
© 2024 Michael C. Orr, Douglas Chesters, Paul H. Williams, Thomas J. Wood, Qingsong Zhou, Silas Bossert, Trevor Sless, Natapot Warrit, Pierre Rasmont, Guillaume Ghisbain, Mira Boustani, A’rong Luo, Yuan Feng, Ze-Qing Niu, Chao-Dong Zhu.
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:
Orr MC, Chesters D, Williams PH, Wood TJ, Zhou Q, Bossert S, Sless T, Warrit N, Rasmont P, Ghisbain G, Boustani M, Luo A’rong, Feng Y, Niu Z-Q, Zhu C-D (2024) Integrative taxonomy of a new species of a bumble bee-mimicking brood parasitic bee, Tetralonioidella mimetica (Hymenoptera, Apoidea, Apidae), investigated through phylogenomics. Journal of Hymenoptera Research 97: 755-780. https://doi.org/10.3897/jhr.97.129470
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A new species of bumble bee-mimicking brood parasitic bee, Tetralonioidella mimetica Orr & Zhu, sp. nov., is described from China. The systematic placement of this species was initially challenging but was resolved using a combination of phylogenomic and COI barcode analyses, which strongly support the new species as a member of the genus Tetralonioidella Strand. Interestingly, the new species mimics the color pattern of both a bumble bee (Bombus Latreille), and its host Habropoda Smith species, a mimicry format previously unknown for bees. A review of the other Tetralonioidella species revealed three additional bee mimics, including two further likely model-host-brood parasite mimicry complexes. To our knowledge, these represent the first documented three-tiered mimetic systems in bees. Several additional taxonomic actions recently became necessary in these and related taxa: Tetralonioidella meghalayensis Dohling & Dey, 2024 is synonymized syn. nov. with Habropoda radoszkowskii (Dalla Torre, 1896) and Varthemapistra Engel, stat. rev. is again synonymized with Habrophorula Lieftinck. Our results also highlight issues with the generic classification of the tribe Melectini as currently used, as Melecta Latreille was found paraphyletic in relation to the remaining melectine genera. As a first step toward resolving this issue, we return the Melecta subgenus Eupavlovskia Popov, stat. rev. to genus level and discuss the ongoing systematic uncertainties regarding melectine taxonomy.
Anthophila, brood parasite, Melectini, mimicry, Nomadinae, taxonomy
The inner workings of mimicry have long fascinated scientists, but relatively few systems have been studied in detail in insects. Special focus has targeted Lepidoptera such as the genus Heliconius Kluk, 1780 and its relatives (
The tribe Melectini contains >200 species of obligately brood parasitic bees worldwide, with especially high species and generic diversity in Asia (
Interestingly, the distribution of Tetralonioidella is concordant with the Asian-Oceanic hotspot of Habropoda Smith, 1854 species richness, and also corresponds closely to the range of Elaphropoda Lieftinck, 1966, which together encompass the known hosts of these brood parasites (
Suggested host associations of Tetralonioidella (=T.). Elaphropoda = E., Habropoda = H. Note that one species may use multiple hosts as seen in other Melectini (
Host | Host author | Parasite | Parasite author | Evidence | Source |
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H. sutepensis | (Cockerell, 1929) | T. habropodae | (Cockerell, 1929) | Co-flight |
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H. christineae | Dubitzky, 2007 | T. heinzi | Dubitzky, 2007 | Elev., phenology, dist. |
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H. bucconis | (Friese, 1911) | T. himalayana | (Bingham, 1897) | Distribution |
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E. erratica | (Lieftinck, 1944) | T. insidiosa | (Lieftinck, 1944) | Co-flight |
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E. impatiens | (Lieftinck, 1944) | T. vulpecula | (Lieftinck, 1944) | Co-flight |
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H. xizangensis | Wu, 1979 | T. himalayana | (Bingham, 1897) | Co-flight, abund., dist. | This study |
H. mimetica | Cockerell, 1929 | T. mimetica | Orr & Zhu, sp. nov. | Co-flight, abund., dist. | This study |
Although Tetralonioidella was originally described as a genus over a century ago (
As part of an initial, systematic treatment of the group we here describe a new mimetic species, Tetralonioidella mimetica Orr & Zhu, sp. nov., and confirm its generic placement based on morphological and molecular analyses, the latter of which is also used to preliminarily investigate the relationships of the genera within Melectini. The remaining Tetralonioidella are then reviewed for additional mimics and potential hosts and mimicry models are discussed for these species. Where relevant, these are assigned to bumble bee color patterns to enable comparison with their geographic ranges. Finally, we briefly consider the phenomenon of mimicry among other Asian bees, with special focus on the Chinese fauna.
A total of seven specimens (three females, four males) of the new species were examined. All specimens directly examined are deposited in the Institute of Zoology, Chinese Academy of Sciences, Beijing (
The terminology used largely follows that of
We constructed a backbone phylogenetic hypothesis using character-rich Ultra-Conserved Element (UCE) data, which was subsequently expanded with placement of additional species represented by COI barcodes only. For UCEs, nine species of Melectini were acquired from
Specimens sequenced. Columns given include published ID number, tribe, genus, species, and the source via which it can be queried. In the source, “here” refers to this paper and in such cases provides the pre-upload voucher code of the sample. GB refers to uploads on GenBank, and BOLD refers to uploads on the Barcode of Life Database.
ID | Tribe | Species | Source |
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CCDB-15281 | Anthophorini | Pachymelus peringueyi | BOLD |
BEECC863-09 | Melectini | Melecta alexanderi | BOLD |
BBHYG927-10 | Melectini | Melecta pacifica | BOLD |
BEECC859-09 | Melectini | Melecta separata | BOLD |
BEECC754-09 | Melectini | Melecta thoracica | BOLD |
BEECB476-07 | Melectini | Brachymelecta californica | BOLD |
BEECC854-09 | Melectini | Zacosmia maculata | BOLD |
KJ839671 | Melectini | Melecta albifrons | NCBI |
KJ839507 | Melectini | Melecta luctuosa | NCBI |
HM401245 | Melectini | Thyreus orbatus | NCBI |
EX037 | Melectini | Melecta albifrons |
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EX036 | Melectini | Melecta italica cf. |
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BLX881 | Melectini | Melecta thoracica |
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EX042 | Melectini | Tetralonioidella pendleburyi |
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EX044 | Melectini | Thyreomelecta sibirica |
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EX088 | Melectini | Thyreus delumbatus |
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EX090 | Melectini | Thyreus quinquefasciatus |
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EX029 | Melectini | Brachymelecta californica |
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D0863 | Melectini | Eupavlovskia obscura | Here: MGPCC007-21 (BOLD) |
D08632 | Melectini | Eupavlovskia obscura | Here: MGPCC008-21 (BOLD) |
IOZ(E)2148081 | Melectini | Tetralonioidella mimetica | Here: MGPCC191-24 (BOLD) |
IOZ(E)2148051 | Melectini | Tetralonioidella wuae | Here: MGPCC192-24 (BOLD) |
BSRUAA6806 | Melectini | Thyreus callurus | Here: PQ074116 (GB) |
BSRUAA6807 | Melectini | Thyreus centrimaculata | Here: PQ074117 (GB) |
BSRUAA6787 | Melectini | Thyreus ceylonicus | Here: PQ074119 (GB) |
BSRUAA6780 | Melectini | Thyreus himalayensis | Here: PQ074118 (GB) |
BSRUAA6801 | Melectini | Thyreus cyathiger cf. | Here: PQ074120 (GB) |
We used the aligned UCE matrix of
We compared the likelihood of this result with other hypotheses to evaluate the generic associations and validity of the taxa included here (with our tree considered hypothesis 1, to see whether alternative scenarios were significantly better supported than our reconstruction when they were imposed via constraints). First, we tested whether Melecta was monophyletic without the tentative subgenus Melecta (Eupavlovskia) Popov, 1955 (hypothesis 2: Melecta monophyletic). We then tested whether (Eupavlovskia) also belonged within the genus Melecta as considered by
The backbone phylogeny provided generally reliable results for the groups of interest in Melectini (Fig.
Topological tests provided definitive insight into the placement of Eupavlovskia, strongly supporting it as separate from Melecta (Suppl. material
The placement of the new species was initially difficult because it exhibited characteristics of both Tetralonioidella and Eupavlovskia. The exceptionally long forewing marginal cell supports the placement of this species in Tetralonioidella, following
Superfamily Apoidea
Family Apidae
Subfamily Nomadinae
Tribe Melectini
Melecta, sensu
Diagnosis is best made in the male sex, as there are no known characters present in the female sex which allow unambiguous separation from all other melectine genera. Generally robust, moderately large bees, 12.5–16 mm in length. Mesosoma covered with long and dense pubescence, this particularly evident on the dorsal surface where it covers and obscures the pronotal tubercules, scutum, scutellum, and scutellar spines. Marginal cell of forewing short, three times as long as broad, only slightly exceeding the third submarginal cell. Labrum almost square, widest basally, surface slightly concave, anterior border entire, little upturned, with rounded side edges, basal tubercles only weakly projecting but generally large. Hind tibia of the male strongly broadened and expanded at its apex, with strong ventroapical process extending laterally beyond the base of the tibial spurs. Inner hind tibial spur noticeably longer than the outer, gently and variably curved in different directions, appearing weakly undulate. Hind basitarsis of the male strongly to moderately broadened in its apical half to two-thirds. Male antennae without rhinarial pits on their posterior faces (sensu Lieftinck). T7 of male subtruncate, apex clothed with appressed tomentum. S7 very slender, with narrow, widely divergent arms and bilobed apex, the lobes fringed with strong bristles; S8 with well-developed ridges in apical half, apex itself with tufts of long feathery hairs.
Due to the thickly hairy mesosoma, Eupavlovskia can appear superficially quite similar to Tetralonioidella, but they may be separated by a short marginal cell (the most common character state for melectine bees) that only extends slightly beyond the apex of the third submarginal cell, the marginal cell itself being clearly shorter in maximum length than the length of the three submarginal cells combined; it is also shorter than the distance between its apex and the apex of the forewing. In Tetralonioidella the marginal cell is much longer, exceeding the third marginal cell and only slightly shorter than the length of the three submarginal cells combined; it is longer than the distance between its apex and the apex of the forewing. From other Eastern Hemisphere melectine bees, Eupavlovskia is separated by the scutellum, which is not flattened into a plate that overhangs the declivity of the propodeum and by the presence of arolia (with a plate-like scutellum and without arolia in Thyreus), by the three submarginal cells (two submarginal cells in Sinomelecta Baker, 1997), by the length of T1, which is dorsally shorter than T2 and the presence of arolia (T1 longer than to scarcely shorter than T2 dorsally and with arolia absent or nearly so in Afromelecta Lieftinck, 1972 and Thyreomelecta), from all Melecta or currently recognized Melecta subgenera by the combination of the long and dense mesosomal pilosity, the shape of the male legs, the absence of rhinaria on the antennal segments, and the structure of the male S7–8.
From Spain in the west across the Western Palearctic to Central Asia (Uzbekistan, Bukhara;
Eupavlovskia, Paracrocisa and Pseudomelecta Radoszkowski, 1865 were separated from Melecta at the generic level by
Tetralonioidella mimetica Orr & Zhu, 2023: holotype (IOZ(E)2148141): male: male, holotype: China, Sichuan Province, Wenchuan City, Yingxiu County, 900 m, 1983.8.3, coll. Zhang Huaicheng. Verbatim: 四川汶川映秀 900m // 1983, 8.3张怀成 // IOZ(E)2148141. Translation: Sichuan Province, Wenchuan City, Yingxiu, 900m // 1983.8.3 Zhang Huaicheng // IOZ(E)2148141. Coordinates from Google Earth retroactive georeferencing: 31.05, 103.49.
The forewing marginal cell is clearly longer than the distance from its apex to the apex of the forewing, and this character separates it from other melectine genera. Additionally, both sexes are immediately recognizable from nearly all other Melectini by color, specifically the yellow scutellar hair followed by a largely black metasoma tipped with reddened terga and hairs. Among melectines, it may still be confused with Tetralonioidella tricolor, from which both sexes can be distinguished by a transverse black stripe of hair on the scutum, and in males additionally by the unmodified hindleg of T. tricolor compared to the enlarged, flattened hindbasitarsus of the new species.
In the key of
Male: pubescence and integumental color: See Figs
Males smaller, body size 12–13mm.
Head : Galea only slightly longer than height of eye, shiny throughout, with minute single-sized punctures, tip angularly pointed but sides rounded to tip. Mandible unmodified, with weak but distinct inferior blade running parallel to main blade. Labrum shiny but somewhat craggy, large punctures indistinct from various angles, rim shallowly but obviously, roundly indented medially, with distinct row of hairs along rim though not forming dense brush. Lacking facial maculations. Clypeus strongly protuberant, by about half max eye width. Clypeus shiny medially near rim, tessellate elsewhere, with distinct rounding outward above shiny rim, with irregular but dense pitting throughout. Cheek immediately slanted inward from rear of compound eye. Antennal F1-2 roughly equal length, slightly longer than subsequent flagellomeres. Ocelli nearly linear, medial ocellus only slightly lower than lateral ocelli. Integumental surface near ocelli shiny, strongly pitted below, slightly tessellate and sparsely, minutely pitted between lateral ocelli and compound eye.
Mesosoma : Intertegular distance (at rear) averaging 3.58 mm based on four specimens (3.5, 3.6, 3.5, 3.7). Wings only very slightly darkened, hairy within cells along fore edge and decreasingly so posteriorly, apical papillae strongly apparent. Scutum, scutellum, metanotum typically obscured by dense, plumose hair. Below, integument densely punctured, interspaces weakly tessellate. Tegula translucent, medium brown, somewhat orange. Scutellar spines large, strongly pointed and directed posteriorly and slightly laterally, still obvious through dense hairs although eclipsed by them. Legs largely unmodified, save for hindleg: tibia in profile increasing in width from unmodified base to tip that is over twice its initial width, vaguely triangular overall, not flattened, broadest apically. Hindbasitarsus similarly narrow proximally, though flattened and broadened to tip like a paddle. Basitibial plate absent.
Metasoma : T1-2 longer from above, about equal, with T3 at most roughly 2/3 of either. Terga covered in small hairs, largely simple medially but increasingly plumose laterally and apically, largely plumose by T4. Terga weakly tessellate between punctures, with weak reflections; usually apparent through appressed setae for T1-2 and often T3. Tergal rims unmodified, of similar opacity, color to rest. Male T6 unmodified. Male T7 in profile gradually thinning to tip with slight abrupt dip medially; without medial longitudinal carina, but covered in dense hairs beyond base; lacking lateral projections or flanges; tip from above bilobed with medial indentation similar in size to each of the lobes, with rounded tip. Male S6 unmodified and lacking distinctive hair patches, very slightly and gradually shallowed medially along rim. Male S7 overall initially appearing disconnected medially given weak medial scleritization contrasting with stronger tan integumental color laterally; with distinct lateral corners nearing 90°, but rounded; with strong subapical hair tufts directed laterally; tip broadly bifid with wide pointed tips, lateral to broad apical, V-shaped emargination. Male S8 shield-like, laterally gradually declivous in latter half though maintaining overall rounded outline until near tip, there projected forward and narrowing to tip, sharply pointed medially with long hairs arising from below. Male genital capsule with outer corners, where gonocoxite tips curve inward to gonostylus, lacking any flange, instead marked by narrowing toward tip, though again slightly expanded terminally, largely without hairs. Interior projection between gonostylus and penis valves also narrowed to tip, but entirely covered in long, plumose hairs largely obscuring form.
Female: highly similar to males overall, differing as given: Pubescence and integumental color: See Fig.
Females slightly larger, in part due to more tapering, elongate metasoma, body size 13–14mm, largest nearing 16mm.
Head : labrum more narrowly and abruptly indented medially, forming clearer corners.
Mesosoma : Intertegular distance (at rear) similar, averaging 3.63mm based on three specimens (3.8, 3.5, 3.6). Legs unmodified, hind tibia only slightly expanded apically, with widest point clearly before tip.
Metasoma : Overall shape roughly similar but tapering to more distinct point terminally. T1-5 visible from above, T6 typically only visible for pygidial plate, itself triangular near base, coming to narrow tip with near-parallel sides. Sterna largely unmodified, last visible sterna narrowed and curled upward, elongate, forming support for very long sting.
This species is recorded from Sichuan (four sites), Guangzhou (one site), and Hunan (one site) at elevations of 700 m, 700–900 m, 900 m, 1150–1200 m, 1270 m, and 1300 m (relatively low for Sichuan bumble bees,
This bee has been collected from July 24 through August 18.
Habropoda mimetica Cockerell, 1927 is the most likely host, based on similarities in known distribution (mid-elevation ringing Sichuan depression and adjacent similar habitat), phenological matching, and elevational similarity. This association was especially evident at the Baishuihe National Nature Reserve in Sichuan, where H. mimetica was exceedingly common, while other species of Habropoda were rarer (Habropoda omeiensis Wu, 1979, Habropoda sinensis Alfken, 1937). It may be that this species targets multiple hosts, but among these H. mimetica is almost certainly utilized.
No floral data are available for this species. Brood parasites are generally considered relatively generalist, given that they need not collect pollen for their offspring, but tracking specific resources may benefit a brood parasite in finding specialized hosts.
The name “mimetica” is given to reference its mimicry and also its likely host, Habropoda mimetica. This name derives from the Ancient Greek adjective mimetikos (that which imitates), and is in the feminine singular nominative form.
All are paratypes except for the holotype: IOZ(E)2148141: male, holotype: China, Sichuan Province, Wenchuan City, Yingxiu City, 900 m, 1983.8.3, coll. Zhang Huaicheng; IOZ(E)2142171: male: China, Chongqing City, Wanzhou District, Wangerbao National Nature Reserve, 1300 m, 1993.8.15, Song Shimei; IOZ(E)2148161: male, genitalia pulled: China, Guizhou Province, Tongren City, Shiqian County, Jinxing village, 700 m, 1988.7.24, coll. Yang Longlong; IOZ(E)2148151: male: China, Hunan Province, Xiangxi Tujia and Miao Autonomous Prefecture, Yongshun County, Muhe Forest farm, 700–900 m, 1988.8.8, coll. Yang Longlong; IOZ(E)2148081: female, EU1 COI voucher: China, Sichuan Province, Baishuihe National Nature Reserve, 31°15'56"N, 103°50'02"E. 2018.8.17, coll. Feng Yuan; IOZ(E)2148071: female: Sichuan province, Chengdu City, Pengzhou City, Xiaoyudong Town, Longcaogou (Baishuihe National Nature Reserve), 31°13'39"N, 103°45'05"E, 1150–1200 m, 2018-VIII-18; IOZ(E)2148061: female: Sichuan Province, Wenchuan City, Yingxiu County, 900 m,1983.8.3, coll. Zhang Huaicheng.
The new species was originally set aside by Yan-Ru Wu as a member of Eupavlovskia, later identified as a possible Tetralonioidella by MCO and ZN and then hypothesized to be Eupavlovskia again by PR.
Bumble bees seem the most likely model in this system. Given the complexity of the color patterns detailed herein, it seems unlikely that they have arisen and been maintained by chance, or by aposematism as might be the case for some all-black, red-tailed species (
If we accept that H. mimetica is the host or one of the hosts of T. mimetica, and that Bombus are the likely models in this system, then this species represents a three-tiered mimicry complex, comprised of the model bumble bee(s), the host H. mimetica, and the brood parasite T. mimetica. In this case, there are many potential bumble bee models, but the most likely models are species including Bombus breviceps Smith, 1852 and Bombus trifasciatus Smith, 1852, based on coloration, distribution, and commonness (
Further investigation of other Tetralonioidella species revealed two additional potential examples of three-tiered mimicry (Fig.
Hypothesized examples of three-tiered mimicry systems in bees. Given are the model (top), likely host (middle), and brood parasite (bottom). System 1: likely model A Bombus lepidus worker (IOZ(E)1429818); likely host B Habropoda xizangensis male (IOZ(E)2051720); and brood parasite C Tetralonioidella himalayana male (IOZ(E)2148111). System 2: likely model D Bombus friseanus worker (IOZ(E)1429817); likely host E Habropoda mimetica female (IOZ(E)2148091); and brood parasites F Tetralonioidella tricolor male (IOZ(E)2148051) G Tetralonioidella mimetica female (IOZ(E)2148071). Note that multiple species may act as models or hosts.
The species of Tetralonioidella with yellow scutal hair and largely black metasomas (e.g., T. pendleburyi, etc.) may also represent mimics of other bees such as Xylocopa appendiculata Smith, 1852, which can be quite common (although more so in or near human habitats). However, this will require further study, as it is an exceedingly common color pattern across various groups of bees, and it is unclear whether this is a result of mimicry (although suggested as such in
The present study highlights the need to solidify and expand our understanding of melectine systematics and taxonomy. As an important first step, we provide a framework and method for integrating data types, which, going forward, will enable quicker and more efficient phylogenomic works on this and other understudied groups for which phylogenomic data are lacking but where many barcodes are available. This is especially important in developing countries where many new species are expected to be present, but where the capacity for phylogenomic sequencing and analysis remains limited (
There are some additional, more recent taxonomic actions which also require further treatment. This year, Dohling and Day (2024) described Tetralonioidella meghalayensis Dohling & Day, 2024 from four female specimens from north-eastern India (Meghalaya state), but the holotype specimen that they imaged is a pollen-collecting bee based upon its visible scopa on the hind legs (see their figs 1, 3). It cannot be Tetralonioidella, as all these brood parasitic species lack scopae. From the character examination (including wing venation, with the 1st and 2nd recurrent veins meeting the 2nd and 3rd submarginal cross veins, respectively) and myriad other features, it clearly belongs to the anthophorine genus Habropoda. Further comparison with
Additionally, the taxonomy of the relatively closely related subfamily Anthophorinae is also in need of clarification. Per
The three-tiered mimicry systems described here raise several interesting new questions that require further investigation. First and foremost, are the selective pressures that resulted in brood parasitic mimics being exerted by predators, or by the hosts themselves? The case for predation driving mimicry has long been established, but it may be initially unclear how hosts could also select for color patterns. In the case of Anthophorinae, they are highly visual bees known to actively come and visually inspect people in the field (
Of interest is also that all confirmed melectine mimics fall within Tetralonioidella. Though almost all melectine bees are associated strictly with Anthophorinae, and therefore, they might benefit from resembling their often violent hosts as described above, no other melectine species conclusively exhibit such coloration. However,
The habitats of the brood parasites, their hosts, and likely Bombus models generally seem to align, especially given that most have been collected together, but this may be due to limited distributional records and collecting effort. The distribution of Tetraloinioidella mimetica, its likely host, and potential Bombus models are interesting from the perspective of color pattern matching. The new species T. mimetica differs slightly from its likely host (H. mimetica) in that the former does not have the transverse black scutum hair stripe of the latter, nor does it have yellow hair on any of the first several terga (Fig.
The interplay of various factors on the quality of mimicry is also a topic worthy of further consideration, as it could help to explain distributional mismatches or some cases of imperfect mimicry. Tetralonioidella are generally medium-sized bees, so under the theory that larger prey animals must better resemble the protected species they imitate (
The bees of Asia present a unique and interesting opportunity for the further exploration of mimicry in bees. For example, the carpenter bees of the subgenus Bombioxylocopa Maa, 1939 all appear to be mimics, and numerous other species have epithets referring to their color patterns resembling bumble bees, such as Amegilla bombiomorpha Wu, 1983 (similar to T. tricolor). However, questions remain as to whether some color forms indicate mimicry or simple aposematism. For instance, there are many Hymenoptera and other insects (potential mimics) that exhibit an all-black form with a red tail, which for aculeate Hymenoptera would clearly indicate the location of the sting. Simple color forms such as this are considerably more difficult to link to mimicry than are the more complex types exhibited by bees such as T. mimetica, given the number of coordinated changes necessary for the latter to evolve, and more work is clearly necessary to further explore these phenomena.
MCO was supported by the National Science Foundation of China’s International Young Scholars Program (31850410464) and The National Science Fund for Distinguished Young Scholars (No. 31625024), and partially by the Chinese Academy of Sciences President’s International Fellowship Initiative (PIFI) (2018PB0003, 2020PB0142, 2024PVC0046). CDZ’s lab is supported by grants from the Key Laboratory of the Zoological Systematics and Evolution of the Chinese Academy of Sciences (grant number 2008DP173354) and the Key Program of the National Natural Science Foundation of China (Grant No. 32330013).
Laurence Packer and Petr Bogusch are thanked for their input on the paper, including photographs from the latter of several species, especially Tetralonioidella tricolor. Yiwei Lu and Mwinzi Duncan Kioko are thanked for logistics support. Author contributions: MCO, PHW, TJW, & CDZ conceived the study; MCO, TJW, SB, TS, NW, PR, GG, MB, AL, FY, & ZN provided specimens and/or data; MCO, PR, AL, & CDZ secured funding; MCO, DC, QZ, & SB conducted analyses; MCO, DC, & SB visualized results; MCO, DC, PHW, & TJW wrote the initial draft; all authors discussed or directly commented on drafts; all authors read and approved the paper.
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