Nesting Biology
Most species of Megachile use existing cavities in which they construct
their nests. Various types of tubular cavities serve as nest
sites. They include dead plant stalks, rolled leaves, spaces
among rocks, termite tunnels and burrows in soil. Old beetle
burrows in wood and holes in masonry are popular sites. The width
of the cavity selected depends on the size of the bee, but most
species use diameters of 10 mm to 20 mm. Several species
accept artificial nests of bamboo canes, bored pieces of wood
and drinking straws. A few excavate their own burrows in the
soil, some use existing holes in the ground, while a number use
either location. In contrast, some species of mason bees build
an exposed nest attached to a wall or branch.
Possibly the first recorded reference to the nesting biology of Megachile
was by Réaumur’s (1742) cited by Kirby (1802) and Jean-Henri
Fabre (1915). Fabre’s accounts of his own detailed observations
on the biology of Megachile (1914, 1915) and other insects
in southern France have fascinated naturalists for years. His
books awakened my own curiosity in wild bees. Fabre gave the
first detailed account of the use of trapnests to study the nesting
biology of these bees. During the same period Ferton (1896, 1897)
studied Megachile species in southern France and on Corsica.
Despite the bees’ economic and ecological importance and the relative
ease of study, the biology of few species has been studied in
detail. This is unfortunate as the readiness of many Megachile
species to utilize trapnests and their fondness for suburban
gardens enables the amateur entomologist to study their nesting
biology with relative ease. Apparently the difficulties of identification
have discouraged such investigations. Detailed studies have been
conducted on M. brevis (Michener 1953) and M. umatillensis
(Bohart & Youssef 1972) in U.S.A. and Krombein (1967) collected
information on several species there. The present author studied
the holarctic species, M. centuncularis in southern
England (Raw 1988). The most detailed studies on neotropical
species are those on M. concinna, M. zaptlana, M. rufipennis
and M. lanata in Jamaica (Jayasingh & Freeman
1980, Raw 1984b, 1985). With hundreds of publications on its
biology, physiology and management, by far the best-known species
is M.. rotundata, the principal pollinator
of alfalfa.
Almost all leafcutter bees construct their nests by a similar method.
A female selects a suitable tubular cavity where she constructs
a series of cells end to end. For each cell she first cuts oval
pieces of green leaves which she uses to form a cup which is the
base of the cell. Next she lines the sides of the cavity with
a number of layers of oval pieces. She then provisions the cell
with food and lays an egg on its surface. Finally she cuts several
leaf discs to seal the cell. The food the mother places in each
cell is a paste of pollen and honey. The cell closure is positioned
at a short distance from the food mass to allow space for the
larva to grow. M. pugnata omits the cell linings
and intercellular partitions. Rarely M. centuncularis
omits the cell lining, the only construction being the leaf discs
placed to separate the cells (Markowsky 1933). Many mason bees
use mud and resin and others chewed leaves to build the partitions
or even the entire cell.
Generally several cells are constructed head to tail in the tunnel.
The females of some species secrete lactones, esters and hydrocarbons
in the Dufour's gland which are added to the provisions, but it
is not known if they are germicides or nutritional supplements
(Williams et al 1986). Megachile is much more efficient
than most herbivorous insects in consuming the stored food. The
growing larva assimilates 54-58% of the energy and 90% of the
nitrogen in real growth (Waldauer 1968, Wightman & Rogers
1978, cited in Roubik 1989: 151 & 282).
Having consumed the food reserve, the final instar larva defaecates
and spins a cocoon. In warmer climates and in regions with longer
summers, development from egg to emergence of the adult can be
completed in about a month. In univoltine species the final larval
instar, called the prepupa, hibernates or aestivates and development
is completed shortly before emergence from the nest.
Females of M. rotundata prefer trap-nests used by the previous
generation to new tunnels. Apparently they detect aromas secreted
left by the previous occupants rather than nest residues (Parker
et al 1976). A nesting female of M. centuncularis
recognizes her own nest as different from those of her conspecifics
(Raw 1992).
Among temperate species a female will normally live up to a month
and produce up to 30 eggs. In warm weather she builds and provisions
about one cell per day. Little detailed information is available
on the detailed nesting behaviour of tropical species.
In many species the female is larger than the male and occupies a
larger cell provided with a greater quantity of food. In a nest
containing both sexes the mother bee commonly lays female eggs
in the inner cells and males in the outer ones (Raw & O'Toole
1979). Many species are protandrous, with the adult males emerging
a week before the females.
As the cells lie head to tail in a narrow tunnel there is a question
of how a bee towards the back of the nest can emerge when siblings
block the exit. Presumably all bees nesting in tunnels behave
as do M. centuncularis, Osmia rufa L, O. ceorulescens
(L) and O. leiana (Kirby) (Raw 1972, 1988). When a bee
towards the back awakens it chews through the partition and gains
access to the cell in front. If that cell is occupied, the bee
nips the occupant. When the bee in front starts to move the bee
behind remains still. Activity of a bee is the signal to the
bee behind to stop biting. When an emerging bee encounters a
dead bee in the cell in front it bites the corpse and, as there
is no response, it chews its way out pushing the remains behind
it.
Mortality
The female of a nesting bee provides the entire food supply for her offspring
so she looks for a secure place to locate the nest. Nevertheless,
this rich supply of food and the defenceless offspring suffer
the attacks from numerous organisms. Predators and other organisms
which kill the developmental stages of solitary bees are of three
sorts. Cuckoo bees and some wasps and beetles kill the egg and
eat the stored food. The parasitoid wasps, Leucospis, Melittobia,
Monodontomerus and Tetrasticus and bee-flies attack
later, killing the growing larva, the pupa or even the adult before
the host can leave the cell. A third group, which includes dermestid
and tenebrioid beetles and flies consume the stored food and apparently
incidentally kill the bee larva.
Many organisms have been recorded attacking Megachile. The
bees are Coelioxys, Dioxys and Stelis. Wasps
include Leucospidae (Leucospis), Eulophidae (Melittobia
and Tetrasticus), Torymidae (Monodontomerus),
Pteromalidae (Pteromalus, Dibrachys and Phaeacra),
Sapygidae (Huarpea and Sapyga), Mutillidae (Dasymutilla),
Chrysididae (Chrysis) and Ichneumonidae (Aritranis
and Sphaeropthalma). Beetles are Dermestidae (Anthrenus,
Trogoderma), Cleridae (Trichodes), Meloidae (Lytta
and Nemognatha, Ptinidae (Ptinus) and Tenebrionidae
(Tribolium, Aprostocetus, Phyllobeanus, Cryptolestes
and Tenebrioides). Flies are Bombyliidae (Anthrax)
and Tachinidae (Aritranis, Dibrachys and Phaenaera).
Two moths have been found consuming the food in the cell (Plodia
interpunctella (Huebner) - the Indian meal moth and Vitula
edmandsae Rogonot - the dried fruit moth). Mites include
Chaetodactylidae (Chaetodactylus), Saproglyphidae (Vidia)
and Suidasiidae (Tortonia).
The most common cuckoo bees attacking Megachile are members
of the genus tip of the Coelioxys The abdomen of the female
of Coelioxys is pointed with which, as first reported by
Ferton, she pierces the wall or the cap of a cell of the bee to
deposit an egg (illustrated in Roubik 1989: 167). The egg hatches
quickly and generally before the host's egg. The first larval
instar is agile and bears long sickle-like mandibles (Graenicher
1905, 1927, Baker 1971). The larva searches for the host bee’s
egg which it punctures with the mandibles and sucks dry. The
second instar larva has normal mandibles and consumes the stored
food of pollen mixed with honey which was provided for the host.
The wasp, Sapyga pumila behaves like Coelioxys (Torchio
1972). The adult female pierces the cap of the cell with her
ovipositor to lay her egg. Likewise, with its long sharp mandibles,
the first instar larva destroys the bee’s egg (and any other wasp’s
egg which may be present) before consuming the stored food. The
eggs of clerid and meloid beetles are laid on flowers. The first
instar larva of the beetle attaches itself to a foraging bee and
is carried back to the nest (Eves & Johansen 1974).
Several management techniques have been developed to diminish the
attacks of predators on species of Megachile of economic
importance as pollinators (Eves & Johansen 1974, Torchio 1972).
The emergence trap utilizes the tendency for predators to emerge
from the cells before their hosts. There are two types, both
employing ultra-violet light which attracts the insects. The
lamp is placed above a tray containing insecticide or oil or is
located inside a box bearing small tubular openings with glue.
In both, the predatory insect is trapped and killed before the
host bee emerges. The method is said to be efficient in controlling
predators in trap-nests that are brought indoors to overwinter.
As the insects which attack the offspring of nesting bees do not construct
their own nests they spend the night in safe cavities. A second
control technique is the use of night station traps which are
similar to trapnests provided for the host bees, but of a smaller
diameter to prevent the bees' entry. However, this technique
must adversely affect the populations of small species of cavity
nesting bees.
Chalkbrood fungus (Ascosphaera species) attack the young stages
of Megachile. In particular A. aggregata specifically
attacks Megachilidae and is a serious cause of death of M. rotundata
(Vandenberg & Stephen 1982). Several studies on Megachile
and related taxa which construct nests comprising linear series
of cells have revealed high rates of unidentified mortality of
the developing brood (Danks 1971, Jayasingh & Freeman 1980,
1984b, 1985, 1988). When an adult female emerges from her natal
nest, she must pass through all the cells in front of her and
thus come into contact with the contents of all the cells including
those in which her siblings died. This is an ideal situation
for a pathogen to pass to the next generation.
Various organisms attack adult Megachile, particularly when
they are on flowers. These include conopid flies (Physocephala
and Megaselia), crab-spiders (Thomisidae) and Strepsiptera.