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Body Form, Tagmata, and Sex Differences

Body Form, Tagmata, and Sex Differences

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9. Body Form, Tagmata, and Sex Differences



prises the second and following metasomal segments, and

for an ant with a two-segmented petiole, the gaster is the

third and following metasomal segments. Thus if one

hopes for a uniform terminology throughout the Aculeata or the Hymenoptera, in which homologous structures have the same names, “gaster” is unsatisfactory, because a particular numbered gastral segment may be

numbered differently in related taxa.

The sexes in bees are often quite different from one another, and in the keys the sexes are sometimes treated separately. Most males have 13 antennal segments or antennomeres, as they are often called, but there are only 12 in

some Euryglossinae (some species of Euryglossina), some

Ammobatini (Pasites, Melanempis, and Parammobatodes), some Ammobatoidini (Holcopasites), some Biastini (some species of Biastes), some Halictini (some

species of Thrinchostoma), and some Augochlorini

(Chlerogas); only 11 or 12 in Systropha (Rophitinae); and



43



14 in Uromonia s. str. (Meganomiinae). Females have 12

antennal segments, although there are only 11 in some

species of Euryglossina (Euryglossinae). Males usually

have seven exposed metasomal terga; females have six.

Sometimes the apical terga are retracted beneath the

preapical ones, so that female Halictinae, for example,

usually appear to have only five terga, and in some male

bees, such as Protosmia, T7 can be seen only with difficulty, because it is largely or wholly retracted and must be

dissected out if one wishes to examine it. Females have

stings, and males have sclerotized genitalia, but both are

usually retracted, and in some females the sting is rudimentary. The sting and associated structures are reduced

and not (or weakly) functional in many Andrenidae;

more reduced, not at all useful for stinging, in the

Meliponini; and maximally reduced, compared to all

other bees, in the Dioxyini. For sting reduction see Figure 27-1.



10. Structures and Anatomical Terminology of Adults

The illustrations and text in this section provide the

names for the external parts of adult bees. Many structures are simply labeled in the illustrations. Those that do

not appear in the figures, or for which discussion or explanation is needed, are treated in the text, with the preferred terms appearing in boldface type at the places

where they are defined or explained. The hope is that the

illustrations plus the text will encourage uniformity of usage in future work.

The emphasis is on features that vary among kinds of

bees; very few terms are explained or illustrated for structures that do not provide useful diagnostic characters in

one or more bee taxa.

The morphology of the honey bee (Apis mellifera Linnaeus), which has been much studied, serves as a background for persons interested in the Apiformes as a

whole. Snodgrass (1956) provided a valuable account of

Apis morphology as well as a terminology for structures

(derived from his earlier studies) that has served as a basis

for subsequent works. Several authors studying non-Apis

bees have given, in varying degrees of detail, their own accounts of external morphology, using the terminology of

Snodgrass except where there were reasons for deviating,

and establishing additional terminologies as needed.

Such studies are those by Michener (1944) on Anthophora; Urban (1967a) on Thygater; Camargo, Kerr,

and Lopes (1967) on Melipona; Eickwort (1969a) on

Pseudaugochlora; Gerber and Akre (1969) on Megachile;

Pesenko (1983) on Nomioides; and Brooks (1988) on An-



thophora. There is no need to repeat here the details of

morphology presented in some of these works, but the

terminology to be used in subsequent parts of this book

requires some explanation. It is that of Michener (1944),

modified in various ways.

Many of the structures referred to in the systematic

parts of this book (Sections 33 to 121) are labeled in Figures 10-1 to 10-15. These figures are mostly diagrammatic, intended to illustrate a maximum number of bee

structures; only a few are based on particular bee species

or genera. Terms that seem obvious from a perusal of these

figures are not further described here. For fuller accounts,

see Michener (1944) and the other morphological studies listed above.

The head. Figures 10-1 and 10-2 illustrate the major

structures of the head. For names of mandibular structures I usually follow Michener and Fraser (1978); see

Figure 10-2. For simplicity I often refer to preapical teeth

on the upper margin instead of teeth of the pollex. The

pollex is the upper margin of the mandible, above the acetabular groove, commonly ending in the preapical tooth

or teeth; the rutellum is the rest of the distal part of the

mandible, below the pollex, usually forming the major or

lower apical mandibular tooth. A few bees (Lithurgus,

many Xylocopinae) have a lower preapical tooth, i.e., a

rutellar tooth, below the main apex (Figs. 78-3a, 90-9a,

b). Other terms used herein are the condylar ridge, which

arises near the mandibular condyle (at the lower or posterior basal angle of the mandible) and extends toward the



Figure 10-1. Diagrams of a

bee’s head, showing major

structures.

a, Anterior view; b, Posterior

view. From Michener, McGinley, and Danforth, 1994.



a



b



44



10. Structures and Anatomical Terminology of Adults



45



a



Figure 10-2. a, Diagrammatic

lateral view of a bee’s head; b,

Outer surface of mandible.

b



From Michener, McGinley, and

Danforth, 1994.



apex of the mandible, and the outer ridge, which is the

next ridge above the condylar ridge on the outer surface

of the mandible.

The terminology for parts of the labrum is often confused by the use, in the genus Andrena, of the term process

for the basal elevated plate. The term process is misleading because this plate does not project, as one expects of

a process. In other bees, e.g., the Panurginae (see Ruz,

1986), the same structure is called the basal area of the

labrum. Use of the word “process” in the sense of basal

area is further confusing because in some bees, especially

the Halictidae, there is an entirely different process on the

apex of the labrum, here called the apical process of the

labrum.

For descriptive purposes the face is often divided into

ill-defined areas, as indicated by dotted lines in Figure 101. These are the two paraocular areas, the supraclypeal

area, the frons or supra-antennal area, and the vertex.

The malar area (or malar space) is between the eye and

the mandible; its length is the shortest distance from the

eye to the mandible (Fig. 10-2); the width of this area is

the width of the base of the mandible.

The foveae of the face (Fig. 10-1) (and of the sides of

T2, Fig. 10-12) are depressions, usually black in color and

therefore conspicuous when the ground color is pale.

Those of the face are paired, one in each paraocular area,

and lie largely or entirely above the level of the antennal

bases, sometimes extending up onto the vertex between

the ocelli and the upper ends of the compound eyes. The

foveae, better developed in females than in males, may be

punctiform or narrow, hairless grooves (e.g., in many Hylaeinae and Euryglossinae) or broad, slightly depressed

areas, those in most Andreninae finely hairy. Schuberth

and Schönitzer (1993) have given an anatomical account

of the facial foveae of various taxa, showing that the epidermis beneath foveal cuticle consists of secretory cells.

Presumably, the foveae are evaporative surfaces, but the



products and their functions are unknown. In many bees

that lack distinct facial foveae there are nonetheless areas

of differentiated cuticle, presumably homologous to

foveae. Such areas are usually more sparsely and less

coarsely punctate than adjacent areas, and frequently differ in surface microstructure and in color (usually black).

Thus every intergradation exists between complete lack

of foveae (as in Megachilidae and most Halictinae) and

distinct, depressed foveae. In keys and descriptions I do

not indicate the presence of foveae unless the area is sufficiently depressed that at least one margin is distinct and

the other indicated by a sharp change in texture or color.

The antennae arise from antennal sockets, sometimes

called alveoli. For simplicity, the terms antennal or flagellar segments are used instead of flagellomeres. Nearly

all bees have a subantennal suture extending from each

antennal socket down to the epistomal suture. If the antennae arise close to the clypeus, there may be no subantennal sutures, and in some bees (most Andrenidae) there

are two subantennal sutures below each antenna, defining a subantennal area. These sutures are easily seen as

dark lines if the integument is pale but are often hard to

see if it is black, and may be invisible if it is both black and

coarsely punctate. The epistomal suture defines the upper limits of the clypeus. A longitudinal carina immediately mesal to the antennal base occurs in some bees. Typically, it is most elevated just above the antennal base and

often forms a lamella that partially overlaps the antennal

base (Fig. 10-3a). Such carinae are often called interantennal or interalveolar carinae. Such terms, however, suggest carinae extending from one antennal base to the

other. The term juxtantennal carinae, proposed by Michener and Griswold (1994a), is therefore preferable for

these structures.

In some bees (e.g., Augochlora) the anterior tentorial

arms seem to have migrated downward, carrying with

them the epistomal suture, which therefore becomes an-



46



THE BEES OF THE WORLD



a

b



c



Figure 10-3. Diagrams of heads and antenna. a, Diagrammatic



nal distance (or length of subantennal suture if it is straight); (7),



frontal view of a bee’s head, showing structures not illustrated in



length of eye; (8) interantennal or interalveolar distance; (9) intero-



Figure 10-1. (No known bee has both paraocular lobes and juxtan-



cellar distance; (10) ocellocular distance; (11) antennocellar or



tennal carinae.) b, Diagrammatic frontal view of a bee’s head,



alveolocellar distance; (12) antennocular or alveolocular distance;



showing how the following measurements are made: (1) length of



(13) clypeocular distance; (14) length of malar area. c, Antenna of



head (or face); (2) width of head; (3) length of clypeus; (4) lower in-



female bee. From Michener, McGinley, and Danforth, 1994.



terocular distance; (5) upper interocular distance; (6) clypeoanten-



gled or lobed down into the clypeus on each side. The resultant lobe of the paraocular area into the clypeus is

called the paraocular lobe (Fig. 10-3a).

The term orbit is often used for the eye margin, inner

orbit for the frontal or facial margin, and outer orbit for

the genal margin. An expression like “eyes converging below” is ordinarily exactly equivalent to “inner orbits converging below.” Imaginary lines tangent to the upper or

lower extremities of both eyes, as seen in a frontal view of

the head, are sometimes useful in indicating the positions

of the ocelli or of the clypeal apex. Such lines are called

the upper and lower ocular tangents.

Descriptions of bees often include measurements or,

more often, statements of relative dimensions of various

structures, especially on the head. Figure 10-3b indicates

how certain measurements should be made. Looking

down on the vertex, one can use the postocular tangent,

the imaginary line tangent to the posterior convexities of

both eyes. The ocelloccipital distance is between a posterior ocellus and the point where the vertex curves or angles down onto the posterior surface of the head, i.e., usually the preoccipital ridge. Several of these terms can be

reversed, e.g., ocellocular has the same meaning as oculocellar.

The genal area is the region behind the eye and in front

of the preoccipital ridge. The ridge surrounding the concave posterior surface of the head above and laterally is

called the preoccipital ridge. A carina sometimes found

on this ridge is the preoccipital carina. It can be dorsal

(behind the vertex only), lateral (behind the eye only), or

complete (both dorsal and lateral). According to Silveira

(1995a), the dorsal part of the preoccipital ridge in some

Exomalopsini is actually on the posterior surface of the

head, and a new transverse ridge (or carina), the postocellar ridge (or carina), is present just behind the ocelli.

The proboscidial fossa (Fig. 10-1) is the large, deep



groove on the underside of the head into which the proboscis folds. The fossa is margined laterally by the hypostomal carina, the anterior end of which bends laterad

behind the mandibular base. The underside of the head,

lateral to the hypostomal carina and behind the mandibular base, is the hypostomal area, or, according to Eickwort (1969b), the postgena; this area is not the entire hypostoma because the latter includes also the walls and roof

of the proboscidial fossa. The paramandibular process is

an anterior projection of the hypostoma that approaches,

butts against, or is fused with the lateral part of the

clypeus, and in the latter case provides sclerotic closure of

the mandibular socket. Bristles or hairs arising from a

ridge on the paramandibular process and sometimes continuing laterally on the lateral extremity of the hypostomal carina constitute the subgenal coronet of some Andrena species.

A disproportionate number of characters used in the

higher classification of bees are based on proboscidial features. Most sclerites of the proboscis are clear from Figure 10-4, but some explanation is needed. Sometimes,

the length of the proboscis is expressed as the point that

it reaches under the body of the bee when retracted,

which means when folded, at rest, or, as often expressed,

in repose. The proboscis consists of three “segments.” The

basal one, containing the maxillary cardines (sing.

cardo), extends backward in repose from their articulations to the cranium. The middle one, containing the

maxillary stipites (sing. stipes), prepalpal parts of the

galeae, and the labial prementum, extends forward in repose from the apices of the cardines. The distal segment,

containing the galeal blades or postpalpal parts of the

galeae, the labial palpi, and the paraglossae and glossa,

extends again backward from the apex of the middle segment. In repose the apices of these third-segment parts

may not reach out of the proboscidial fossa, or may ex-



10. Structures and Anatomical Terminology of Adults



47



b



a



c

d



Figure 10-4. Diagrams of proboscides of bees. a, Spread

proboscis of a long-tongued

bee; b, Maxilla of the same; c,

Labium of a short-tongued, in

this case colletid, bee, showing

portions of maxillary cardines

at the base; d, Maxilla of the

same. From Michener, McGinley, and Danforth, 1994.



tend back on the ventral surface of the body to or even beyond the apex of the metasoma. The proboscis is extended, for example to probe a flower for nectar, by unfolding these segments. It can project downward from the

bee’s head, or forward. For descriptive purposes I consider

it to project downward, but some other authors make the

other decision. What I consider the anterior surface of the

proboscis, they call the dorsal surface.

Proboscidial structures are illustrated and labeled not

only in Figure 10-4 but also in Figures 19-1 and 19-2. The

terminology of the parts of the glossa is extensive and is

indicated in Figures 19-3 and 19-4.

The galeal blade, i.e., the postpalpal part of the galea,

is divided by a longitudinal galeal rib into (1) a thin, anterior, hairless part, its sclerotized surfaces compressed together (Roig-Alsina and Michener, 1993, fig. 15), called

by J. Plant (manuscript, 1991) the galeal velum; and (2)

a posterior part that supports hairs, and whose inner and

outer surfaces are largely separated. In S-T bees the apices

of these parts are commonly separated by a notch near the



apex of the galeal blade. The galeal rib, which often bears

a series of hairs, appears as a strengthening element, principally in L-T bees. The prepalpal part of the galea is the

subgalea.

There has been confusion about the naming of the

basal sclerites of the labium. The main labial sclerite,

which is in the middle “segment” of the labium and to

which the labial palpi are attached, is regularly called the

prementum. Basal to the prementum, and supported in

the membrane between the apical parts of the maxillary

cardines, are one or two sclerites that can be called the

postmentum. In most Hymenoptera there is only one

such sclerite. In many bees there is a sclerite immediately

basal to the prementum that tapers basally and is thus

more or less triangular. This is what I call the mentum

(Michener, 1944, 1985a), following Snodgrass, who,

however, in 1956 called it the postmentum. Plant and

Paulus (1987) consider it the distal part of the postmentum, the basal part being the lorum. In some bees the

mentum is partly (as in some Andrena and Panurgus) or



48



THE BEES OF THE WORLD



wholly (as in Ctenocolletes, some Andrena) membranous

(Michener, 1985a), but can be identified by its position

and shape. In other bees, such as the Halictidae, the membranous area is smaller, not so distinctively shaped, and

may represent either the reduced mentum or merely a

membrane between the prementum and the postmentum. Basal to the mentum, in most bees, is the lorum. It

was called submentum by Michener (1944), but Winston

(1979) and others pointed out that because there is only

one basal labial sclerite in other Hymenoptera, the lorum

should be considered a new structure, special to the bees,

not homologous to the submentum of some other insect

orders. Plant and Paulus (1987), however, as indicated

above, regarded it as the basal part of the postmentum.

For convenience, I term the labial sclerites, starting

basally, to be lorum, mentum, and prementum. When

the lorum expands as a weak sclerotization occupying

space between the cardines, I call it the loral apron.

When, as in most Halictidae, the mentum is partly or

wholly membranous and doubtfully recognizable, I

nonetheless tentatively regard the single basal labial sclerite as the lorum and loral apron, not as a fusion product

of lorum plus mentum. These sclerites are fused in some

L-T bees, but this is not as common or so evident as was

indicated by Plant and Paulus (1987), who illustrated

quite distinct sclerites as fused in some cases.

The basal part of the proboscis, i.e., the part that attaches to the head, is called the labiomaxillary tube (stippled in Fig. 21-2). Its skeletal parts are the cardines, which

are strong rods in the wall of the tube, and it is further

strengthened by flexible strips called the conjunctival

thickenings. (Additional longitudinal thickenings on the

anterior surface are of maxillary lacinial origin in the Halictidae.) For illustrations, see Figures 21-2b and 59-1. In

many bees the lower end of the conjunctival thickening

is separated as a small sclerite, the suspensorium of the

prementum, that connects the thickening to a lateral

notch in the prementum. In the posterior surface of the

labiomaxillary tube, the lorum varies greatly, as described

above and by Michener (1985a). It may be a rather weak,

flat loral apron, i.e., a thickening or sclerotization of the

posterior wall of the labiomaxillary tube occupying most

of the space between the cardines. It may be more limited

in area but elevated around the connection to the mentum, or reduced to a strongly sclerotized, V-shaped structure connected medially to the mentum (Fig. 10-4a). In

the last two alternatives, when the labium is retracted, the

base of the mentum and median (apical) part of the lorum together commonly project, forming a lobe extending posteriorly from the labiomaxillary tube. This lobe is

the proboscidial lobe (Fig. 21-2a). In some bees, when

the proboscis is retracted, this lobe projects upward and

into the postoccipital pouch, a large pit below the foramen magnum (Roig-Alsina and Michener, 1993).

The thorax. The bee thorax (Fig. 10-5) is a compact

structure consisting of sclerites of the pro-, meso- and

metathoracic segments, which bear the legs and wings,

and the first true abdominal segment, termed the

propodeum. The prothorax is represented primarily by

the large pronotum, which extends ventrally at each side

as a process that meets or nearly meets its fellow behind

the fore coxae. The propleura and prosternum are in



front of this lateroventral extension of the pronotum. The

pronotal lobe is a useful landmark. The dorsolateral angle of the pronotum is in front of and somewhat mesad

from the pronotal lobe; often there is a ridge, carina, or

lamella connecting the lobe to the dorsolateral angle.

Sometimes a ridge, carina, or elevated zone extends between the two dorsolateral angles along the posterior

margin of the pronotum; this is the pronotal collar. Another ridge or carina may extend directly downward from

the dorsolateral angle toward the front coxa.

In dorsal view, the mesothorax can be divided into four

distinct sclerites: the scutum, the scutellum, and the

paired axillae. The suture between each axilla and the

scutellum is the axillar suture; other sutures in this area

are easily recognized by name, e.g., scutoscutellar. Laterally, the mesothorax is represented by the mesepisternum, sometimes referred to as the mesopleuron. The

mesepisternum is sometimes divided by the nearly vertical episternal groove, formerly called the pre-episternal

groove. The episternal groove may extend down, after

meeting the anterior end of the horizontal scrobal

groove, onto the lower anterior part of the mesepisternum, as in most Colletinae and Halictinae (Fig. 20-5b);

it may curve posteriorly in an arc that merges indistinguishably with the scrobal groove as in Andrena and many

Apinae (Fig. 20-5c); or it may be absent (Fig. 20-5a) or so

short as not to reach the scrobal groove, as in the

Megachilinae. The scrobe is a small pit on the scrobal

groove in front of the meso-metepisternal suture. The

area above the scrobal groove and behind the episternal

groove is often more convex and shiny than adjacent areas; it is the hypoepimeral area (not labeled in Fig. 10-5).

The depressed and largely hidden anterior margin of the

mesepisternum is the prepectus, according to Brothers

(1975).

The lateral (as distinguished from ventral) part of the

mesepisternum is divisible into an anterior-facing surface

and a lateral-facing surface. The angle between these surfaces varies from gradually rounded (Fig. 20-5a) through

sharply angular to carinate and even lamellate. To avoid

expressions like “angle between anterior and lateral surfaces of mesepisternum,” Michener and Griswold

(1994a) introduced the term omaulus for this angle. For

sphecoid wasps, “omaulus” is used only if the angle is

carinate; for bees the term is broadened and used even if

the angle is rounded, so that one can record “omaulus

rounded” or “omaulus carinate,” or even “omaulus lamellate.” The omaulus, which is not shown in Figure 10-5, is

anterior to the episternal groove when the groove is present, and the preomaular area is anterior to the omaulus,

i.e., it is the anterior, forward-facing surface of the

mesepisternum.

Dorsally, the metathorax consists of a sclerite, the

metanotum, which is obliquely divided at each side by

the transmetanotal suture. The metepisternum (or

metapleuron) forms the lateral surface of the metathorax.

The wing bases are located above the upper margins of

the mesepisternum and the metepisternum.

The middle and hind coxae of bees seem superficially

to be shifted posteriorly so that the middle leg appears to

arise from the lower end of the metepisternum and the

hind leg from the propodeum (Fig. 10-5a). Of course this



10. Structures and Anatomical Terminology of Adults



49



a



Figure 10-5. Diagrams of a

bee’s thorax. a, Lateral view; b,

Dorsal view. (The tegula is

omitted in a and the left side of

b. The propodeal triangle is indicated by dotted lines in b.)

b



From Michener, McGinley, and

Danforth, 1994.



is not true, as careful examination shows; nevertheless the

middle coxa, which in many bees is expanded upward to

form a vertically elongated cylinder (see Michener,

1981b), displaces or is partly hidden beneath the lower

part of the metepisternum.

The form and subdivisions of the propodeum are important systematically. Many bees have a pair of impressed lines on the propodeum (dotted in Fig. 10-5b),

beginning near its anterior dorsolateral parts and extending downward and posteromedially and nearly meeting

in or above the propodeal pit, a median depression of

the lower posterior surface. These lines, together with

the anterior dorsal margin of the propodeum, enclose the

propodeal triangle. Morphologically, this triangle is

the metapostnotum (Brothers, 1976). The shape of the

propodeum as seen in profile is quite independent of the

triangle. The whole propodeum may be vertical or nearly

so, dropping from the posterior margin of the metanotum (Fig. 20-5a, c). In this case it is termed declivous.

But there is frequently a more or less horizontal or sloping basal region (Fig. 20-5b), sometimes separated by a

sharp line or carina from the declivous posterior surface,

as shown in Figure 10-5a. The horizontal part is called the



basal zone or basal area of the propodeum, sometimes

called the enclosure when it is set off or enclosed by carinae; usually its sculpturing is distinctive, e.g., with striae

radiating from its base. The basal area may be part of the

propodeal triangle, or may extend beyond the triangle, at

least laterally. Sometimes the basal area, as recognized by

its sculpturing, is vertical like the rest of the posterior

propodeal surface. The term “basal area” is applicable

even if no sharp line separates it from the vertical surface

and even if it is slanting or vertical rather than horizontal. In some bees the two surfaces are continuously

rounded, one onto the other in a broad, curving surface;

in that case the term “basal area” is not definable unless

there is distinctive surface sculpturing. In a few bees the

triangle is reduced in size, its lateral margins meeting and

continuing posteriorly as a single line (sometimes not recognizable) to the propodeal pit. In other cases (e.g., in

some Xylocopa) the reduction seems to have continued

until there is no triangle, but only a median longitudinal

line extending to the pit. In other bees, when the triangle

is not recognizable, it is because the lines that demarcate

it are weak or absent.

The wings. Wings are illustrated, and the veins la-



50



THE BEES OF THE WORLD



Figure 10-6. Diagram of the

wings of a bee, showing the

vein terminology of Michener

(1944). From Michener, McGinley, and Danforth, 1994.



beled, in Figure 10- 6, using a modified Comstock and

Needham system. Wings are described as though spread,

so that the direction toward the costal margin (where the

stigma is in the forewing) is called anterior; toward the

wing apex, distal. To save space, the word stigma is used

in place of pterostigma. Because the homologies of the

veins are not very certain, as well as because of some comparable-looking veins that often are referred to as a group,

yet have very different morphological names, it has

seemed best to continue the use of terms that are morphologically noncommittal for certain cells and veins

much used in taxonomy. The names of cells and certain

noncommittal names for veins are shown in Figure 10-7.

Table 10-1 gives the equivalents, in Comstock-Needham

terms, of these names.

Of special importance are three veins that all look like

crossveins: the second abscissa of Rs, first r-m, and second

r-m, to use the Comstock and Needham system. These

veins help to define the submarginal cells, which are usu-



ally either three or two in number. When there are only

two submarginal cells, one sometimes does not know

whether the missing vein is the second abscissa of Rs or

the first r-m; both losses can apparently occur, and both

result in two submarginal cells, as illustrated by Peters

(1969). Hyleoides (Colletidae) (Sec. 47) illustrates the impossibility of knowing which vein is lost. Hylaeinae have

two submarginal cells; in most genera the first is much

longer than the second, suggesting that the first is really

the fusion product of the first and second, but in Hyleoides

the reverse seems to be true. Expression is greatly simplified by using similar terminology for all three veins. In the

past they have been called first, second, and third transverse cubital veins. These veins, however, have nothing to

do with the cubitus; in fact the cubitus is in a very different part of the wing. I prefer to call them first, second,

and third submarginal crossveins (1st, 2nd, 3rd in Fig.

10-7). This terminology agrees approximately with that

of Diniz (1963), which was proposed for the same reason.



Figure 10-7. Diagram of the

wings of a bee, showing the terminology of cells and morphologically noncommital terms for

certain veins. (The notations

1st, 2nd, and 3rd refer to the

submarginal crossveins.) From

Michener, McGinley, and Danforth, 1994.



10. Structures and Anatomical Terminology of Adults



Table 10-1. Morphologically Noncommittal Terms

for Certain Forewing Cells and Veins and Their

Equivalents in Comstock-Needham Terminology.

Noncommittal terms

marginal

first submarginal

second submarginal

third submarginal



Comstock-Needham terms

Forewing Cells

2nd R1

1st R1

1st Rs

2nd Rs



Forewing Veins

basal

M

first recurrent

1st m-cu

second recurrent

2nd m-cu

first submarginal crossvein

2nd abscissa of Rs

second submarginal crossvein

1st r-m

third submarginal crossvein

2nd r-m

prestigma

1st abscissa of R1

anterior margin of marginal cell 2nd abscissa of R1

posterior margin of marginal

r and Rs

cell

posterior margins of submarRsϩM, 2nd and

ginal cells

following abscissae

of M



It has the drawback that the first is not technically a

crossvein, but is thought to be a transverse section of a

longitudinal vein, Rs.

Louis (1973) reviewed prior alar terminologies and

proposed a new nomenclature for veins, attempting to

avoid considerations of homology and phylogeny. He

called the submarginal crossveins the first to third RM, or

radiomedial veins. I believe that the less technical expression, submarginal crossveins, gives a clearer indication of

what these veins are really like and of their relation to the

submarginal cells.

Figure 10-8 shows the stigma, marginal cell, and

nearby structures, with lines to show how measurements

that are used in the keys and by various authors are to be

made. The width of the prestigma is measured to the

costal margin of the wing, and the expression is thus a

misnomer, for it is more than the width of the prestigma



Figure 10-8. Forewing of Ceratina rupestris Holmberg, showing

how certain measurements are made, as follows: (a) length of

stigma; (b) length of costal edge of marginal cell or of margin of cell

on the costa or wing margin (not a useful measurement in a wing

like this, in which the cell diverges gradually from the wing margin);

(c) length of marginal cell beyond stigma; (d) length of marginal cell;

(e) length of free part of marginal cell; (f ) length of prestigma; (g)

width of prestigma (to wing margin); (h) width of stigma. The breaks

in the wing veins are the alar fenestrae.



51



proper. The length of the costal edge (or margin) of the

marginal cell or of the marginal cell on the costa is repeatedly used. This is the measurement from the apex of

the stigma distad to the apex of the cell, or, if the cell is

truncate, to the point where the cell diverges abruptly

from the wing margin. As suggested by Figure 10-8, it is

a poor measurement to use for wings in which the marginal cell bends gradually away from the wing margin. A

common usage is basal vein for the first abscissa of vein

M in the forewing (Fig. 10-7).

The jugal lobe and vannal lobe of the hind wing are

both measured from the wing base to the apices of the

lobes. Thus, on Figures 10-6 and 10-7 one might say that

the jugal lobe is about two-thirds as long as the vannal

lobe. The terminology of the veins and cells of the posterior parts of the wings varies; the word “vannal” is sometimes replaced by “anal,” and the abbreviation V by A.

Thus the vannal lobe is sometimes called the anal lobe.

The alar fenestrae are small, clear areas occupying specific sites in various veins (Fig. 10-8). Flexion lines, often

faintly visible in the wing membrane, cross veins at these

fenestrae.

The legs. Some authorities advocate a system for identifying parts of legs that assumes that all legs are pulled

out laterally at right angles to the long axis of the body.

Although I appreciate the logic of that system, I here follow the more traditional system in which the legs are considered to be in their normal positions. Thus, the corbicula of corbiculate Apidae is on the outer, not the anterior,

surface of the hind tibia, and the two hind tibial spurs are

outer and inner, not anterior and posterior. Some additional positional terminology is indicated by numbers in

Figure 10-9.

The tibial spurs are the movable inferior apical spurs

on the tibiae; there is one spur (part of the strigilis) on the

front tibia, one on the middle tibia, and in nearly all bees

two on the hind tibia. The tibial spines (Fig. 10-9) are immovable, sharp, superior apical projections, usually small

in size, often blunt or minute, found in some bees. There

are none, one, two, or rarely three spines per tibia; often

they are mere angles. The tibial spur of the front leg consists of a main axis or malus, and a thin platelike scraper

or velum directed toward the main axis of the leg; the

velum usually does not extend to the apex of the malus.

In some Apinae a prong or projecting ridge on the anterior side of the malus is termed the anterior velum by

Schönitzer (1986) and Schönitzer and Renner (1980).

The inner hind tibial spur of the hind leg is especially important taxonomically. This spur usually has two toothed

margins. It is the inner one that is commonly elaborated

in various ways. Following custom, I have described this

margin as ciliate if it has slender, almost hairlike projections (usually numerous), although in many cases the appearance is like that of a fine comb. Because the finely serrate or ciliate (or intermediate) condition is common in

Hymenoptera (Gennerich, 1922) and frequently the

same as that of the outer hind tibial spur and of the middle tibial spur, such spurs are often described as simple,

meaning, I suppose, unmodified. It may be, however, either coarsely serrate or pectinate. Again following custom, I have described a spur as pectinate if its inner margin is produced into several long, coarse, often blunt



52



THE BEES OF THE WORLD



projections, even though the number of such projections

is in some cases reduced to only one or two. The sockets

of the hind tibial spurs and their relation to the tibia vary

among taxa and were the subject of a study by Cane

(1979).

The basitibial plate (Figs. 8-9, 10-9) is on the upper or

outer side of the base of the hind tibia of many bees. It is

best developed in females and presumably is important

for support as bees move up or down their burrows in the

soil or tamp the cell surfaces with the pygidial plate. The

importance of the latter function is suggested by the observation that most female bees either have both pygidial

and basitibial plates or lack both. Commonly, the basitibial plate is surrounded by a carina or a sharp line of

some sort and its vestiture (if any) differs from that of adjacent regions, but it may be indicated only by a series of

tubercles, as in many Euryglossinae, or even by a single

tubercle that indicates its apex; in some cases (as in many

Xylocopa) its apex is represented by a structure near the

middle of the tibia instead of more or less near the base.

In some bees (e.g., certain groups of Centris) the basal or

central part of the basitibial plate is sharply elevated above

the rest of the plate surface. Such an area is called the secondary basitibial plate.

Most S-T bees possess a pair of brushes or combs, best

developed in females, on the middle legs. One is on the

underside of the tibia, sometimes also including a brush

on the basitarsus. The other is on the basal part of the femur, sometimes extending onto the trochanter. These

structures are opposable and are used in cleaning or transferring pollen from the ipsilateral (same side) foreleg

(Jander, 1976). They are termed the midtibial brush or

comb and the midfemoral brush or comb (see Sec. 13).

A comb is a single row of bristles, whereas a brush is less

organized.

On the inner surface of the hind tibia of most bees is

an area of variable size covered with hairs of uniform

length, usually blunt, truncate, or briefly bifid. These

hairs, the keirotrichia (Fig. 10-11), appear to serve for

cleaning the wings. In some bees they are replaced by

longer, more ordinary hairs that may function as part of

the scopa in females.

On the hind basitarsus of many female bees is a distal

process that extends beyond the base of the second tarsal

segment. (For simplicity, I use the expression “tarsal segment” instead of tarsomere.) Sometimes this process

bears on its apex a small brush, the penicillus (Fig. 10-9).

(This brush bears no relation to the tibial tuft known as

the penicillum in the Meliponini.)

Between the tarsal claws there is often a protruding,

padlike arolium (Fig. 10-10). Its lower distal surfaces are

almost always dark, often black, a fact that helps to distinguish it from associated pale structures. See Michener

(1944) and Figure 10-10 for details of the structures between the claws. It is likely that comparative study of

these structures would yield new characters of value for

bee phylogeny or systematics, although loss of the

arolium has occurred repeatedly among bees.

The scopae. Female bees have scopae for holding and

transporting pollen; males do not. Exceptions are the Hylaeinae and Euryglossinae in the Colletidae, parasitic and

robber bees in various families, and queens of highly eu-



Figure 10-9. Hind leg of a female bee, hairs omitted except those

that form the penicillus. The numeral 1 indicates the posterior or upper margin of the tibia; 2, the outer surface; and 3, the distal or apical margin. Modified from Michener, McGinley, and Danforth, 1994.



social bees (Meliponinae and Apinae), all of which lack

scopae. In the keys and descriptive comments I often refer to the scopa without reminding the user that scopae

are found only on females. The scopa consists of pollencarrying hairs. These are not usually the hairs and brushes

with which pollen is removed from flowers, but are the

brushes on which pollen is carried back to the nest. Some

pollen may be carried on various parts of the body, but

scopae occur principally on the hind legs (Figs. 6-4, 8-5)

or on the metasomal sterna (Fig. 8-7). In most bees the

scopal hairs are on the hind legs, but in nonparasitic

Megachilidae they are on the metasomal sterna; in some

colletids and halictids they are on both the underside of

the metasoma and on the hind legs. If fringes of scopal

hairs surround a space in which pollen is carried, they are

said to form a corbicula. The best-known corbicula, on

the outer side of the hind tibia of the corbiculate Apidae

(Euglossini, Bombini, Meliponini, Apini), consists not

only of the fringes of hairs but also of the enclosed concave or flat surface (Fig. 10-11). Other corbiculae are on

the undersides of the hind femora of Andreninae, Halictidae, Colletidae, and others, and on the sides of the

propodeum of many species of Andrena, some halictids,

and some colletids.

The metasoma. For simplicity and to save space, as

noted in Section 9, metasomal terga and sterna are referred to as T1, T2, etc., and S1, S2, etc., T1 and S1 constituting the basal segment of the metasoma (Fig. 10-12).

Each metasomal tergum or sternum (except for the anteriormost and the reduced apical ones) consists of a plate

commonly marked by some transverse lines, as follows.

First, across the anterior margin, always completely hidden in the intact metasoma, is the antecostal suture. The

equivalent internal ridge is the site of attachment of longitudinal intersegmental muscles. The very narrow rims

of the tergum and sternum anterior to the antecostal suture are the acrotergite and acrosternite, plus the apodemal margin that expands to form apodemes laterally. Second, nearer to the middle of each plate is another

transverse line, the gradulus. Typically, the surface basal



10. Structures and Anatomical Terminology of Adults



53



b

c



a



d



to the gradulus, i.e., the pregradular area or disc, is

slightly elevated compared to that behind it, rendering

the gradulus a minute step, as shown in Figure 10-12b.

The ends of the tergal graduli, unless bent strongly to the

rear, are usually near the spiracles. If bent strongly to the

rear, the resultant longitudinal lines are called lateral

parts or lateral arms of the graduli or, if carinate, lateral gradular carinae, sometimes elevated to form lateral gradular lamellae or lateral gradular spines. The

graduli—except for their lateral arms, when present—

are often concealed by the preceding terga or sterna on

the intact metasoma but, especially on T2 and S2, are

sometimes exposed or can be exposed easily by slightly extending the metasoma artificially. Third, near the posterior margin of each tergum and sternum is usually another transverse line, the premarginal line, separating the

marginal zone (posterior marginal area of Michener,

1944; apical depression of Timberlake, 1980b) from the

rest of the sclerite (Fig. 10-12). This zone is often depressed but in other cases differs only in sculpturing from

the area basal to it; sometimes the marginal zone is not

differentiated at all. The region between the gradulus and

the premarginal line can be called the disc when a name

is needed for it. Sometimes, e.g., on T2 of Exomalopsis,

the premarginal line is arched far forward, so that the

marginal zone is broad and the disc reduced to a transverse zone. The dorsolateral parts of the tergal discs (between the graduli and the premarginal lines), especially

on T2 to T4, are often somewhat elevated, convex, and

frequently shiny. These dorsolateral convexities frequently accentuate the premarginal lines, which limit the

convexities posteriorly. The narrow posterior margin of

the marginal zone is often recognizably different from the

rest of the zone in sculpture or is elevated; this is termed

simply the margin. Packer (2004a) provided an account

of sternal variation among bees.

T1 differs from other terga because its base is constricted for the narrow connection with the thorax. Its

dorsal, horizontal surface is nonetheless similar to that of



Figure 10-10. Dorsal views of apices of tarsi. a, Andrena mimetica

Cockerell; b, Anthophora edwardsii Cresson; c, Xylocopa orpifex

Smith; d, Anthidium atripes Cresson. Arolia are well developed in a

and b, absent or greatly reduced in c and d. The median, aroliumlike structure in d is membranous, not obvious in dry specimens,

and Anthidium is considered to lack arolia. From Michener, 1944.



succeeding terga, often having a marginal zone, premarginal line, and disc, each easily recognizable. Toward the

base, T1 is strongly declivous. Often there is a transverse

line or carina near the summit of the declivous surface and

more or less separating it from the horizontal surface.

This line or carina may be the gradulus of T1, although I

do not use that term for it. It is the line or carina delimiting the anterior surface or anterior concavity of T1, and

it is well developed in Heriades and certain other small



a



b



Figure 10-11. Hind tibia and basitarsus of a worker of Plebeia

frontalis (Friese). a, Outer surfaces, showing the scopa reduced to

fringes around the smooth and largely hairless tibial corbicula; b, Inner surfaces. From Michener, McGinley, and Danforth, 1994.



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