A new subgenus of Megachile from Borneo with arolia (Hymenoptera, Megachilidae)

2006·0.72 MB·English

by  Baker, Donald B

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PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
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Number 3505, 12 pp., 13 figures, 1 table January 12, 2006
A New Subgenus of Megachile from Borneo with
Arolia (Hymenoptera: Megachilidae)
DONALD B. BAKER1 AND MICHAEL S. ENGEL2-3
CONTENTS
Abstract . 2
Introduction . 2
Systematics . 2
Genus Megachile Latreille . 2
Matangapis, new subgenus . 2
Megachile (.Matangapis) alticola Cameron. 6
Discussion . 8
Acknowledgments . 10
References . 10
Appendix 1: Notes on Two Oriental Megachile “Subgenera” . 11
1 24 Chichester Court, Old Schools Lane, Ewell, Surrey K17 1TP, UK (deceased. May 10, 2004).
2 Division of Invertebrate Zoology, American Museum of Natural History; Division of Entomology (Paleoento-
mology). Natural History Museum, and Department of Ecology & Evolutionary Biology, 1460 Jayhawk Boulevard,
Snow Hall, University of Kansas, Lawrence, Kansas 66045-7523 ([email protected]).
3 To whom all correspondence should be addressed.
Copyright © American Museum of Natural History 2006 ISSN 0003-0082
2 AMERICAN MUSEUM NOVITATES NO. 3505
ABSTRACT
A new subgenus of the bee genus Megachile (Megachilinae: Megachilini) is described from
Mt. Matang in Borneo. Matangapis, new subgenus, is proposed for Megachile alticola Cam¬
eron, and is most noteworthy for the possession of arolia on all pretarsi of both sexes. The
significance of arolia on M. alticola is briefly discussed in the context of the distinctiveness
of the tribes Megachilini and Osmiini. Notes on two Oriental subgenera of Megachile, Orien-
tocressoniella and Neocressoniella, are appended. The following taxonomic changes are pro¬
posed: Orientocressoniella is a new junior synonym of Callomegachile, Neocressoniella is a
new junior synonym of Xanthosarus, Anthophora barbata Fabricius is a new synonym (senior
but suppressed) of M. carbonaria Smith, M. amputatiformis Cockerell is a new junior synonym
of M. saigonensis Cockerell, M. saphira Cockerell is a new junior synonym of M. Ulrica
Nurse. In addition, the following five species are newly transferred to subgenus Chelostomoda:
M. Ulrica, M. lefroma Cameron, M. albolineata Cameron, M. funnelli Cockerell, and M. bou-
gainvillei Cockerell. Lectotypes are newly designated for M. alticola and M. anthracina Smith.
INTRODUCTION or near Mt. Matang but not, apparently, ob¬
tained elsewhere. The species is remarkable
The bee tribe Megachilini is a cosmopol¬
among Megachile most notably for the pos¬
itan and distinctive lineage of long-tongued
session of arolia on all pretarsi of both sexes.
bees and presently comprises three genera—
Indeed, under the more highly split classifi¬
the cleptoparasitic Coelioxys and Radosz-
cations of Megachilini the taxon considered
kowskiana and the free-living species of
herein would be accorded generic rank. Mor¬
Megachile (table 1). The latter genus, which
phological terminology in the descriptions
includes among its members the familiar
generally follows that outlined by Engel
leaf-cutter bees, is remarkably diverse and
(2001).
has in the past been segregated into multiple
genera (e.g., Michener, 1962; Mitchell,
SYSTEMATICS
1980). Indeed, once the relationships of the
various subgroups within Megachile (sensu
Genus Megachile Latreille
Michener, 2000) are established, a multige¬
Matangapis, new subgenus
neric system might be warranted. The phy-
logeny of Megachile is presently under crit¬ Type Species: Megachile alticola Camer¬
ical investigation by Mr. V.H. Gonzalez, and on, 1902.
it is hoped that much light will eventually be Diagnosis: Size small (ca. 8 mm); habitus
shed on the natural classification of the lin¬ compact (figs. 1-2); metasoma in both sexes
eage. At present one of the least understood short, in dried material globular, not or little
faunas for Megachile, and many bees in gen¬ longer than mesosoma; vertex and genae
eral, are those of Asia. Numerous species are very coarsely, reticulately punctate, several
known to exist in this region, some with times more coarsely than mesoscutum (as in
character combinations that may prove to be Chelostomoda)-, female mandible quadriden-
crucial in resolving megachiline evolution. tate, male mandible bidentate; all distitarsi
Unfortunately, the monographic work that is aroliate; distitarsi coarsely setose, apex of
needed as the foundation for such studies has each subangularly excised dorsally, accom¬
not been undertaken. The present brief con¬ modating base of pretarsus; pretarsus with
tribution is designed to provide insight into orbicula well developed and bearing two
the taxonomic placement of one such critical strong setae, a distinct unguifer not observed,
species as well as to provide some taxonomic probably usually invaginated within distitar-
notes toward clarifying the confusing Asiatic sus in dried material; arolium well developed
Megachile fauna. (fig. 5) but weakly sclerotized, consequently
Herein is provided an account of a strange, usually seen crumpled in dried material; a
long-described but apparently little-known, camera distinct; pretarsal ungues (claws) bi¬
Bornean species, Megachile alticola Camer¬ fid in male, simple in female.
on, 1902, obtained by several collectors on Description: Male. Head in frontal aspect
2006 BAKER AND ENGEL: MEGACHILE ALTICOLA FROM BORNEO 3
TABLE 1
Current Hierarchical Classification of Tribe Megachilini
(based on Michener, 2000, and synonymies proposed herein: vide appendix)
—Tribe MEGACHILINI Latreille—
Genus Megachile Latreille subgenus Paracella Michener
subgenus Acentron Mitchell subgenus Parachalicodoma Pasteels
subgenus Amegachile Friese subgenus Platysta Pasteels
subgenus Argyropile Mitchell subgenus Pseudocentron Mitchell
subgenus Austrochile Michener subgenus Pseudomegachile Friese
subgenus Austromegachile Mitchell subgenus Ptilosaroides Mitchell
subgenus Callomegachile Michener subgenus Ptilosarus Mitchell
subgenus Cestella Pasteels subgenus Rhodomegachile Michener
subgenus Chalicodoma Lepeletier de Saint Fargeau subgenus Rhyssomegachile Mitchell
subgenus Chalicodomoides Michener subgenus Sayapis Titus
subgenus t Chalicodomopsis Engel subgenus Schizomegachile Michener
subgenus Chelostomoda Michener subgenus Schrottkyapis Mitchell
subgenus Chelostomoides Robertson subgenus Stellenigris Meunier
subgenus Chrysosarus Mitchell subgenus Stelodides Moure
subgenus Creightonella Cockerell subgenus Stenomegachile Pasteels
subgenus Cressoniella Mitchell subgenus Thaumatosoma Smith
subgenus Cuspidella Pasteels subgenus Trichurochile Mitchell
subgenus Dasymegachile Mitchell subgenus Tylomegachile Moure
subgenus Eumegachile Friese subgenus Xanthosarus Robertson
subgenus Eutricharaea Thomson subgenus Zonomegachile Mitchell
subgenus Gronoceras Cockerell Genus Coelioxys Latreille
subgenus Grosapis Mitchell subgenus Acrocoelioxys Mitchell
subgenus Hackeriapis Cockerell subgenus Allocoelioxys Tkalcu
subgenus Heriadopsis Cockerell subgenus Argcoelioxys Warncke
subgenus Largella Pasteels subgenus Boreocoelioxys Mitchell
subgenus Leptorachis Mitchell subgenus Coelioxys Latreille
subgenus Litomegachile Mitchell subgenus Cyrtocoelioxys Mitchell
subgenus Matangapis Baker & Engel subgenus Glyptocoelioxys Mitchell
subgenus Maximegachile Guiglia & Pasteels subgenus Haplocoelioxys Mitchell
subgenus Megachile Latreille subgenus Liothyrapis Cockerell
subgenus Megachiloides Mitchell subgenus Neocoelioxys Mitchell
subgenus Megella Pasteels subgenus Platycoelioxys Mitchell
subgenus Melanosarus Mitchell subgenus Rhinocoelioxys Mitchell
subgenus Mitchellapis Michener subgenus Synocoelioxys Mitchell
subgenus Moureapis Raw subgenus Torridapis Pasteels
subgenus Neochalicodoma Pasteels subgenus Xerocoelioxys Mitchell
subgenus Neochelynia Schrottky Genus Radoszkowskiana Popov
transverse (fig. 4), width: length (to clypeal clypeus as in most Pseudomegachile). Lower
margin) ratio 1: 0.85. Vertex short, lateral part of gena not structurally modified but
ocelli separated from preoccipital ridge by with long, dense pubescence in hypostomal
less than twice their diameter; preoccipital area. Mandible slender, apical margin
ridge sharply angular dorsally, becoming car¬ oblique, bidentate (fig. 7); surface between
inate laterally. Clypeal margin unmodified, the coarse, elongate punctures glossy; juxta-
straight; surface of clypeus moderately con¬ genal process absent. Antenna moderately
vex, coarsely and densely punctate with long, elongate, reaching in repose to slightly be¬
erect, simple setae arising from punctures, in¬ yond mesosoma; A13 compressed and slight¬
tegument glossy between punctures, apically ly expanded.
(as adjacent parts of paraocular areas) with Mesosoma short, length (from anterior
appressed plumose pubescence (i.e., the margin of mesoscutum to posterior margin of
4 AMERICAN MUSEUM NOVITATES NO. 3505
Figs. 1-2. Photomicrographs of Megachile (Matangapis) alticola Cameron. 1. Female, lateral hab¬
itus. 2. Male, lateral habitus.
scutellum): width (between tegulae) ratio 1: Metafemur and metatibia moderately thick¬
0.96. Pretarsal ungues cleft and arolium pres¬ ened; metatibia slightly longer than metafe¬
ent on all legs; procoxa unmodified, mutic, mur, at its thickest, at about two-thirds of its
anterior surface without cluster of rufescent length, not thicker than metafemur; two cal-
setae, simply, uniformly pubescent; profe¬ caria present; metafemur ventrally with thin
mur, protibia, and protarsus unmodified; pro¬ fringe of long, simple setae; metatibia ven¬
tarsus slender, with short, even, anterior trally with thin fringe of long, coarsely plu¬
fringe and longer, irregular posterior fringe; mose setae; metatarsus without conspicuous
probasitarsus parallel-sided, about three setal fringes.
times as long as broad; speculi absent. Mid¬ Metasomal terga without distinct marginal
legs unmodified, mesotibial calcar present; areas, with marginal fasciae, these short and
mesotarsal fringes as described for forelegs. lateral on Tl, meeting or nearly meeting on
2006 BAKER AND ENGEL: MEGACHILE ALTICOLA FROM BORNEO 5
Figs. 3-4. Photomicrographs of Megachile (Matangapis) alticola Cameron. 3. Female, facial view.
4. Male, facial view.
T4; T1 short, its dorsal surface medially not teeth. Venter with three normally exposed
greatly longer than ocellar diameter, thinly sterna; SI deeply, angularly emarginate, the
clothed with long, erect, pale setae; T2 and two sides of the emargination forming an an¬
T3 weakly punctate; T4 and T5 with weak, gle of about 100°, the base of the emargina¬
postgradular sulci, sulci not fasciate; T6 in tion enclosed by a V-shaped depression
median third with weak transverse ridge not whose apex extends nearly to the base of the
forming a distinct carina, in profile tergum sternum, this depression clothed with fine,
concave before ridge, weakly convex beyond decumbent, golden pubescence, laterally
it, margin without lateral teeth; T7 short, in sternum rather narrowly exposed, broadly
profile its dorsal surface concave, ending in overlapped by ventrolateral extension of Tl,
a distinct lip that in dorsal aspect occupies forming a nearly parallel-sided plate; apically
median third of its width, ventral surface nar¬ angles of emargination rather broadly round¬
row, apical margin simply arcuate, without ed; S2 without special structural characters,
apically truncate, with dense, broad marginal
fascia of white plumose setae; S3 with weak
median longitudinal sulcus, apical margin
broadly convex, disc thinly clothed with ap-
pressed, white, plumose setae, this clothing
becoming denser apically on either side of
median sulcus; S4 weakly sclerotized, short,
rectangular (fig. 8), its apical margin weakly
incised medially, the area raised laterally, en¬
closed by graduli, which are strongly con¬
vergent apically and approach apical margin
at about one-third inward from its lateral an¬
gles, minutely setose; S5 much narrowed me-
sad, its apicolateral lobes with erect, pale se¬
tae arising from a dense, conspicuous cluster
Fig. 5. Scanning electron micrograph of the of black alveoli, inwardly of each apicolater¬
metapretarsus and metadistitarsal apex of Mega¬ al lobe with a broad marginal tract, decres¬
chile (Matangapis) alticola Cameron depicting
cent mesally and not quite reaching midline,
large arolium. Arolia are present on all legs of
of stronger setae arising from pale, well-sep¬
both sexes; the claws of the female (seen here)
are simple, whereas those of males are bifid. The arated alveoli, these setae either inclined or,
small, spheroid objects on the claws are grains of toward the margin, strongly bent, mesally
pollen. (fig. 9); S6 lacking special structural or setal
6 AMERICAN MUSEUM NOVITATES NO. 3505
characters, its apical margin biemarginate, in length one sixth of tergal width; T6 en¬
forming a convex median lobe and lateral tirely covered dorsally with appressed pale
lobes (fig. 10); S8 of simple outline, shaped plumose setae. Sterna without marginal fas¬
as in M. (Hackeriapis) trichognatha Cock¬ ciae; scopa thin, of rather stiff, simple setae,
erell, 1910 (cf. Michener, 1965: 190, his fig. pale except at extreme sides and distally on
654), apically, laterally, with a few very S6, where it is black; S6 without glabrous
weak marginal setae. Genital capsule short marginal areas.
and broad (figs. 12, 13); gonocoxa with Etymology: The new genus-group name
short, subacute, lateral process; gonocoxae is based on the type locality, Gunong Ma-
apically divergent, decurved, decurved por¬ tang, Sarawak combined with Apis (Latin,
tion in lateral aspect triangularly expanded “bee”). The name is feminine.
(fig. 12); penis valves basally widely sepa¬
rated, convergent apically, their apices Megachile (Matangapis) alticola Cameron
roundly expanded. Figures 1-13
Female. Head in frontal aspect transverse
Megachile alticola Cameron, 1902: 118 ($<?).
(fig. 3), width: length (to clypeal margin) ra¬
tio 1:0.83. Vertex short, lateral ocelli sepa¬ Diagnosis: As for the subgenus (vide su¬
rated from preoccipital ridge by little more pra).
than their own diameter; preoccipital ridge Description: Cameron’s (1902) descrip¬
sharply angular dorsally, becoming subcari- tion is inaccurate, quite apart from an obvi¬
nate laterally. Clypeal margin weakly con¬ ous error in his description of the female
cave medially; general surface of clypeus where “scape” on p. 119 should be “scopa”,
nearly plane, coarsely, reticulately punctate, and may be corrected and amplified as fol¬
together with supraclypeal and interantennal lows:
areas clothed with stiff, pale, suberect, api¬ As for the genus with the following addi¬
cally proclinate setae, the proclinate portions tions: Female. Length ca. 8.1 mm; forewing
more or less spiriform. Mandible quadriden- length ca. 5.5 mm. Clypeus (except the
tate (fig. 6), without intercalary cutting edg¬ weakly concave anterior margin narrowly)
es; third tooth weak (as counting from bot¬ densely, strongly, subreticulately punctate;
tom to top), its sides forming an angle of supraclypeal area equally strongly, reticulate¬
about 165°. ly punctate, especially medially and superi¬
Mesosoma short, length (from anterior orly the punctures tending to coalesce lon¬
margin of mesoscutum to posterior margin of gitudinally. Mesoscutum semimatt, regularly
scutellum): width (between tegulae) ratio 1: reticulately, rather shallowly, punctate, ex¬
0.94. Pretarsal ungues (claws) simple, not cept its anterior declivity where the puncta-
cleft; arolium present (fig. 5). Anterior and tion becomes fine and dense; general surface
intermediate legs without special characters. of scutellum similarly punctate; mesepister-
Metafemur of normal proportions, more slen¬ na, except anteriorly, strongly, reticulately to
der than mesofemur; metatibia slightly lon¬ subreticulately punctate, glossy. Basal area of
ger than metafemur, expanded apically, api¬ propodeum finely coriaceous basally, the
cally slightly wider than greatest width of fe¬ sculpture becoming obsolete posteriorly;
mur, its posterior surface entirely clothed posterior face of propodeum, outside the bas¬
with pale keirotrichia; metabasitarsus broad, al area, glossy with well-separated punctures.
subbasally as wide as greatest width of me¬ T2-5 glossy, densely, regularly punctate, the
tatibia, posteriorly entirely and anteriorly in punctures finer than those of mesoscutum;
ventral half clothed with a dense brush of T1 dorsally more finely punctate, especially
black setae. apicad.
Metasomal terga without postgradular sul¬ Modified setae of clypeus and supracly¬
ci and without distinct marginal areas; Tl-2 peal area yellowish; erect setae of paraocular
with weak, inconspicuous, lateral, marginal areas anteriorly and gena white, a conspicu¬
patches of pale stramineous pubescence, T3 ous but narrow tract of denser white pubes¬
with denser, conspicuous, lateral fasciae of cence adapted to lateral epistomal sulci; erect
white pubescence that do not, however, attain setae of vertex black. Pronotum dorsally
2006 BAKER AND ENGEL: MEGACHILE ALTICOLA FROM BORNEO 7
Figs. 6-13. Megachile (Matangapis) alticola Cameron; 6, female; 7-13, male. 6. Female mandible.
. . .
7 Male mandible. 8. Fourth metasomal sternum. 9 Fifth metasomal sternum. 10 Sixth metasomal
. . .
sternum. 11 Eighth metasomal sternum (setae omitted). 12 Genital capsule, lateral aspect. 13 Genital
capsule; left half is dorsal aspect, right half is ventral aspect.
thinly clothed with short, white setae, later¬ setae; marginal areas of T2 and T3 with con¬
ally pronotal lobes with dense white pubes¬ spicuous, sublateral patches of pure white se¬
cence; mesoscutum with short, inconspicu¬ tae, punctiform on T2, forming a short fascia
ous, black setae, scutellum with longer, black on T3 (Cameron’s “the segments are proba¬
setae; mesoscuto-scutellar sulcus with incon¬ bly edged with white pubescence” was ill
spicuous, narrow, fasciae of white setae ad¬ founded); T6 only (not the apical two seg¬
joining axillae; mesepisterna thinly clothed ments) dorsally with depressed gray pubes¬
with white setae, more or less mixed with cence; setae of dorsal surface of metasoma
black dorsally; propodeum laterally with lon¬ otherwise inconspicuous, black. Scopal setae
ger white setae. Setae of legs for greater part long, simple, off-white; no underlying mar¬
black, forming a dense brush on mesobasi- ginal fasciae present.
tarsus, but fine sericeous setae clothing entire Male. Length ca. 7.6 mm; forewing length
posterior surface of metatibia white. Hori¬ ca. 5.1 mm. Clypeus discally glossy, with
zontal surface of T1 thinly clothed with long well-separated punctures. Punctation other¬
white setae, T1 and T2 laterally with white wise generally similar to that of the female,
AMERICAN MUSEUM NOVITATES NO. 3505
but disparity between sizes of punctures treated as a tribe, Osmiini, equivalent to the
more marked, very coarse on vertex, fine on Megachilini, one of the principal characters
metasomal terga. used in distinguishing them has been the
Vestiture generally similar to that of fe¬ presence or absence of arolia, these having
male. Clypeus discally with erect simple se¬ until recently been held to be uniformly ab¬
tae, laterally and apically, as also paraocular sent in the latter tribe. Two megachiline
areas anteriorly, with subdecumbent plumose groups, however, do possess arolia, Heria¬
pubescence; supraclypeal and frontal areas dopsis Cockerell, 1931, with arolia present
also with subdecumbent pubescence but here on the anterior and intermediate pretarsi in
directed vertically; black setae of vertex and both sexes, and Matangapis, with arolia pres¬
of mesonotum longer and more conspicuous. ent on all pretarsi in both sexes. The exis¬
Setae of all tarsi pale fulvous; protarsus with tence of a Bornean megachiline with all tarsi
short (not exceeding probasitarsal width), aroliate was noted, without further details
regular, moderately developed posterior and without naming the species, by Michener
fringe, exceeded by some longer setae of out¬ (2000: 559), who had seen specimens of M.
er surfaces of tarsal segments. Marginal fas¬ alticola when visiting the senior author in
ciae of terga more strongly developed, short, 1995. The arolia in Heriadopsis and Matan¬
lateral, on Tl, progressively longer, attenuate gapis are likely not indicative of a basal
mesally, T2 and T3, not or only narrowly placement for these subgenera within Mega¬
separated medially on T4; T5 and T6 without chilini as their other characters are suggestive
special setal characters, weakly pale-pubes¬ of more derived positions within the Chali-
cent, T5 with scattered, long erect setae. S2 codoma series of subgenera. Instead, it is
with broad marginal fascia, nearly divided more likely that they have reacquired the ar¬
medially, of decumbent white plumose setae; olia otherwise lost within the tribe.
S3 except laterally and narrowly in median Peters (1970: 199)4 recognized that Her¬
line, clothed with fine, decumbent, plumose iadopsis, described in a paper entitled “Her-
pubescence that becomes denser, and direct¬ iadine and related bees from Liberia and the
ed laterally, toward its apex, leaving, how¬ Belgian Congo” (Cockerell, 1931), was in
ever, a small, glabrous median lip (S4 nor¬ fact a megachiline, although he remained un¬
mally concealed); S4-6, S8 depicted in fig¬ certain as to its status vis-a-vis Chalicodoma
ures 8-11. and chose to treat it as of generic rank rather
Type Material: Of three registered syn- than associate it with such other forms as
types in the Natural History Museum, Lon¬ Chelostomoides Robertson, 1901, Hacker-
don, one 6 and two ?, the 6, B.M. Type iapis Cockerell, 1922, and Chelostomoda
Hym.l7a 2057b, is now designated as lec- Michener, 1962, as a subgenus of Chalico¬
totype. The type series was collected by G.E. doma or Megachile s.l. Michener (2000: 559)
Bryant at Matang, Sarawak (3,000 ft), one of treated Heriadopsis as a subgenus of Mega¬
numerous collectors who have visited the lo¬ chile in an essentially retrograde classifica¬
cality. tion that subsumed all nonparasitic mega¬
Additional Material: 2?, 2d, labeled chilines in a single genus. Megachile so con¬
“N.W. Borneo, / 3600 feet / Mt. Matang, stituted is an enormous and morphologically
near / Kuching, Sarawak. / capt. June 1900 / very diverse assemblage, and attempts to
by Dyak collector. / Presented 1900 by / R. date have, in the absence of a fully compre¬
Shelford, M.A.” (Donald & Madge Baker hensive cladistic analysis, embracing not
Collection, Division of Entomology, Univer¬
sity of Kansas Natural History Museum).
4 According to Peters (1970: 203), the distribution of
Heriadopsis striatulus Cockerell, 1931, was possibly
DISCUSSION limited to S. Katanga; Michener (2000) added Zimbab¬
we and Malawi. A long series of both sexes from Zam¬
Where the osmiines, whether including bia is in the University Museum of Oxford and are la¬
beled as: “N.E. Rhodesia: Luwingu to mouth of Cham-
(e.g., Michener, 2000) or not (e.g., Robert¬
bezi River, x [October] 1908, S.A. Neave” and “Lower
son, 1903; Michener, 1941; Peters, 1970;
Kalungwesi Valley, dense forest, 3500 ft, 12-13 ix [Sep¬
Griswold, 1985) the heriadines, have been tember] 1908, S.A. Neave”.
2006 BAKER AND ENGEL: MEGACHILE ALTICOLA FROM BORNEO
only named genus-group entities but also in¬ coarse punctation of the head, but it differs
formally recognized groupings and individ¬ in a number of significant details apart from
ual highly aberrant species, inevitably failed the possession of arolia, including, in both
to yield any universally acceptable scheme of sexes, the much shorter vertex, which in
classification. The fact that there exists a Chelostomoda is three or more times as long
number of elements that would appear to as the ocellar diameter. In the male, it differs
defy classification does not, however, justify in the bidentate, not tridentate, mandible; in
failure to recognize that included under Mega¬ the proportions of basitarsus and tibia6; in the
chile s.l. there are numerous genus-group
edentate apical margin of T6; and in the en¬
taxa that are as distinct as any genera of bees,
tirely different, complex, form of the geni¬
and that, although their exact status may not
talia (cf. Michener, 2000: 536, fig. 82-10m).
yet be clear, most if not all the residuum of
In the female, it differs in the unusual setal
groups and species can safely be associated
characters of the face; in the mandible having
with one or another of those genus-group
four, not five, teeth and no intercalary cutting
taxa. As a practical convenience, aberrant
edge; in T2 and T3 lacking postgradular sulci
taxa could temporarily be classified under the
and distinct marginal areas; in T6 being
names of most-probable relatives by such a
clothed with pale tomentum; and in the ster¬
simple expedient as using an informal group
na lacking apical fasciae.
or species name in combination with that of
Although Heriadopsis and Matangapis are
the putative relative; e.g., Chalicodoma
at present the only apparent exceptions to the
(Pseudomegachile [amputatum-group]) san-
absence of arolia in all Megachilini, their ex¬
dacanum (Cockerell, 1919) or Megachile
\Eutricharaea-Mf.] orientalis Morawitz, istence does to some extent vitiate the dis¬
1895, the square brackets denoting that the tinctness of the tribes Megachilini and Os-
enclosed term was not intended to have any miini (or Osmiini/Heriadini). Michener
nomenclatural standing under the Code. This (2000: 426), in keying the tribes of the Mega-
might prove to be an intermediate solution, chilinae, employed only two other characters
permitting the segregation of these distinct for separating the Megachilini and the Os-
groups and placement of aberrant species miini, one, in practice rather challenging, re¬
while cladistic work is ongoing. lating to the orientation and vestiture of the
Matangapis belongs clearly to Michener’s preaxilla, the other, neither universal nor
(2000) “chalicodomiform” series, although clear-cut, relating to the presence or absence
its habitus is distinctly not chalicodomiform. of integumental metallic coloration. It will be
In his keys to Palearctic and Oriental Mega¬ apparent that any student dealing with unfa¬
chile s.l. (Michener, 2000: 544-545), the miliar and unusual Megachilinae must pay
male runs best to Chelostomoda, the female attention to the presence or absence of arolia
to Pseudomegachile; in his earlier key to in addition to more commonly observed
Australian Chalicodoma, the male again runs
characters. Certainly, to revert to the recog¬
best to Chelostomoda, the female to the en¬
nition of a single tribe Megachilini for the
tirely dissimilar Chalicodomoides. Superfi¬
megachilines, osmiines, and heriadines (e.g.,
cially, except in its short and thick, not slen¬
Michener, 1944) is unwarranted and not to
der-bodied and parallel-sided, habitus, Ma¬
be preferred. Future cladistic and taxonomic
tangapis does resemble Chelostomoda,5 per¬
studies must focus on more rigorously defin¬
haps most strikingly in the exceptionally
ing higher clades among the Megachilinae.
5 Chelostomoda includes, in addition to the species
indicated by Michener (1965: 204), Megachile Ulrica 6 Exceptionally, in Chelostomoda, the protarsus may
Nurse, 1901 [= M. saphira Cameron, 1907, new syno¬ be moderately expanded (e.g., as in M. Ulrica) or all the
nym]; M. lefroma Cameron, 1907; M. albolineata Cam¬ tarsi may be highly modified, the pro- and mesotarsi
eron, 1897; M. funnelli Cockerell, 1907; and M. bou- greatly expanded, the metabasitarsus contracted and ex¬
gainvillei Cockerell, 1911, all newly placed in subgenus panded, the mediotarsal segments very short but pro¬
Chelostomoda herein. There are also several undescribed longed distad on either side in slender digitiform pro¬
species known from Sri Lanka, India, Thailand, the Ma¬ cesses, and the distitarsus elongated and arched, con¬
lay Peninsula, and Indo-China. spicuously longer than the other tarsomeres combined.
10 AMERICAN MUSEUM NOVITATES NO. 3505
ACKNOWLEDGMENTS of bees—XCV. Annals and Magazine of Nat¬
ural History, series 9, 10: 265-269.
We are grateful to Dr. Michael Ohl and Dr. Cockerell, T.D.A. 1927. Some bees, principally
Dorothea Bruckner for their valuable com¬ from Formosa and China. American Museum
mentary on the manuscript. Partial support Novitates 274: 1-16.
for this study was provided by NSF EF- Cockerell, T.D.A. 1931. Heriadine and related
0341724 (to MSE). This is contribution num¬ bees from Liberia and the Belgian Congo (Hy¬
menoptera, Apoidea). Revue de Zoologie et de
ber 3428 of the Division of Entomology,
Botanique Africaines 20(4): 331-341.
Natural History Museum, University of Kan¬
Engel, M.S. 2001. A monograph of the Baltic am¬
sas.
ber bees and evolution of the Apoidea (Hy¬
menoptera). Bulletin of the American Museum
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