As also expected from the evolutionary argument, mating patterns vary greatly from one species to the

next. However, most of the patterns thus far studied fall into one or the other of two broad classes, or
"syndromes" (Hölldobler and Bartz, 1985). In the first, the female-calling syndrome, the females, which
are often wingless and sometimes just fertile workers, do not travel far from the nest. Standing on the
ground or low vegetation, they release sex pheromones to "call" the winged males to them (Figure 3-5).
This pattern is displayed by Amblyopone and Rhytidoponera, which are members of the phylogenetically
primitive subfamily Ponerinae (Haskins, 1978); presumably also by the very primitive Nothomyrmecia
macrops (Hölldobler and Taylor, 1983); at least one pseudomyrmecine, the Neotropical acacia ant
Pseudomyrmex ferrugineus (Janzen, 1967); and the socially parasitic species of the myrmicine genera
Doronomyrmex, Formicoxenus, Harpagoxenus, and Leptothorax (Buschinger, 1968a,b, 1971a,b, 1975b;
see Figure 3-6).

In general, the colonies of female-calling species are typically small at maturity, with 20 to 1,000
workers, and produce relatively few reproductives. So far as known the females mate only once. An
unusual variation on this pattern is followed by the Florida harvester ant Pogonomyrmex badius.
Females gather on the surface of their home nest and are inseminated by males; afterward they fly off
to start new colonies. Van Pelt (1953) thought that the males came from the same nest as the females
with whom they copulate, but S. D. Porter (personal communication) observed that they usually fly for
about a quarter-hour first before settling on a nest different from their own. Porter observed one case in
which a male mated with two females after alightingAs also expected from the evolutionary argument,
mating patterns vary greatly from one species to the next. However, most of the patterns thus far
studied fall into one or the other of two broad classes, or "syndromes" (Hölldobler and Bartz, 1985). In
the first, the female-calling syndrome, the females, which are often wingless and sometimes just fertile
workers, do not travel far from the nest. Standing on the ground or low vegetation, they release sex
pheromones to "call" the winged males to them (Figure 3-5). This pattern is displayed by Amblyopone
and Rhytidoponera, which are members of the phylogenetically primitive subfamily Ponerinae (Haskins,
1978); presumably also by the very primitive Nothomyrmecia macrops (Hölldobler and Taylor, 1983); at
least one pseudomyrmecine, the Neotropical acacia ant Pseudomyrmex ferrugineus (Janzen, 1967); and
the socially parasitic species of the myrmicine genera Doronomyrmex, Formicoxenus, Harpagoxenus,
and Leptothorax (Buschinger, 1968a,b, 1971a,b, 1975b; see Figure 3-6).

In general, the colonies of female-calling species are typically small at maturity, with 20 to 1,000
workers, and produce relatively few reproductives. So far as known the females mate only once. An
unusual variation on this pattern is followed by the Florida harvester ant Pogonomyrmex badius.
Females gather on the surface of their home nest and are inseminated by males; afterward they fly off
to start new colonies. Van Pelt (1953) thought that the males came from the same nest as the females
with whom they copulate, but S. D. Porter (personal communication) observed that they usually fly for
about a quarter-hour first before settling on a nest different from their own. Porter observed one case in
which a male mated with two females after alighting
The second combination of traits during mating is the male-aggregation syndrome. Males from many
colonies gather at specific mating sites, usually prominent features of the landscape such as sunflecked
clearings, forest borders, hilltops, the crowns of trees, and even the tops of tall buildings. Sometimes, as
in some species of Lasius and Solenopsis, the males cruise in large numbers at characteristic heights
above the ground. The females fly into the swarms, often from long distances, in order to mate (see
Figures 3-7 through 3-9 and Plate 2), and afterward they typically disperse widely before shedding their
wings and excavating a nest. The winged queens and males of the fire ant Solenopsis invicta, for
example, fly up to heights of 250 meters or more; 99 percent then descend to the ground within a 2-
kilometer radius of their origin, while a very few travel as far as 10 kilometers. The ability of a single
mature colony to disseminate fertile queens in many directions over long distances is one of the reasons
the fire ant is so difficult to eradicate (Markin et al., 1971). Male-aggregation species typically differ from
those utilizing female calling in two other key respects: the mature colonies are large, containing from
several thousand to over a million workers and producing hundreds to thousands of reproductive adults
yearly, and multiple insemination is common. An unusual reversal of the usual swarming procedure was
recently discovered in some Pheidole species of the southwestern United States: the winged queens
gather in aerial swarms, where they maintain a more or less uniform distance from each other while
attracting males with pheromones. The males fly into the female swarms and mate with individual
females (Hölldobler, unpublished). Swarms of variable composition, some predominantly male and
others predominantly female (occasionally exclusively female), have been reported by Eberhard (1978)
in the coccid-tending formicine Acropyga paramaribensis of northern South America.

Ant species can be classified another way into two broad types. When the males alight on the surface of
the mating site, either in response to female calling or in swarms to compete directly with one another,
they are often typically large and robust in form and possess well-developed mandibles. In contrast,
males that gather in aerial swarms are usually (but not invariably) smaller relative to the queen than are
males of the first type. Also, their mandibles are reduced in size and dentition, sometimes consisting of
nothing more than vestigial lobate or strap-shaped organs. An example of this type is the small
myrmicine Pheidole sitarches of the southwestern United States. Up to 50 males form circular swarms
that hover from a few centimeters to two meters above the surface of woodland clearings. The virgin
queens fly in slow, even circles through the aggregations until mounted in midair by a male, whereupon
the pair cease flying and spiral to the ground together to complete the copulation (Wilson, 1957b).

The swarms of some ant species are among the more dramatic spectacles of the insect world. W. W.
Froggatt (in Wheeler, 1916c) describes the flight of the giant Australian bulldog ant Myrmecia sanguinea
as follows:
References

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