Parelhoenders in Handbook of The Birds of The World 01
Parelhoenders in Handbook of The Birds of The World 01
Parelhoenders in Handbook of The Birds of The World 01
Volume 2
New World Vultures to Guineafowl
• Medium-sized, plump terrestrial birds, with small head, and extensive bare skin on he
head and neck; most species with crest or bony casque.
• 40-72cm
• Afrotropical Region.
• Wide variety of habitats, from dense rain forest to semi-desert.
• 4 genera, 6 species, 19 taxa.
• 1 species threatened; none extinct since 1600
Systematic
The guineafowl have traditionally been considered a subfamily of an expanded version of
Phasianidae, but recent studies based on differences in DNA indicate that family treatment is
appropriate for the group. According to DNA sequences the Numididae diverged from the
Phasianidae lineage some 38 million years ago.
The Family Numididae is endemic to Africa, and its closest relatives are apparently to
be found in Phasianidae and Cracidae. It seems most likely that the guineafowl originated in
the savanna zones of Asia, from a francolin-like phasianid, which probably colonized Africa
some time around the middle to late Miocene. The changes in the distribution patterns of
vegetation over geological time, determined by fluctuations in the climate and also by
geological activity, influenced the taxonomic development of the ancestral guineafowl by
causing a considerable degree of radiation. This, in turn, has led to the various representatives
of the family occupying practically all of the biotopes found in African in the present day.
All four of the genera currently recognized within the guineafowl were clearly
differentiated by the Pleistocene. Of the four, Agelastes is probably the most primitive. Its two
species constitute a superspecies. The little known Black Guineafowl (Agelastes niger) was
formerly awarded a separate genus, Phasidus, but in the early 1960’s a detailed comparison of
this species and the White-breasted Guineafowl (A. meleagrides) led to the conclusion that the
two species were closely related, and should be united in a single genus.
Over the years, the Helmeted Guineafowl (umidida meleagris) has been subdivided
into some 30 subspecies, but at present only nine are generally recognized, due to extensive
intergradation. Two or sometimes three of these races have actually been considered separate
species by some authors, and, while there is only limited support for this approach, there are
three fairly clearly defined subgroups, with the nominate form occupying East Africa, the
galeata group in West Africa, and the mitrata group in southern Africa. As in many similar
cases, it is essentially a matter of personal preference as to whether these groups are treated as
megasubspecies or allospecies but zones of intergradation and the element of caution both
argue in favour of the former.
The Plumed Guineafowl (Guttera plumifera) and the Crested Guineafowl (Guttera
pucherani) form another super-species. Some authors have preferred to recognize three
species in the genus Guttera, dividing the Crested Guineafowl into two, with the promotion of
the race edouardi, but once again extensive interbreeding, both in the wild and in captivity,
has shown that they should be considered conspecific, although they have perhaps moved
some little way towards speciation. About 15 subspecies have been described for the Crested
Guineafowl, but currently only five of these are usually accepted.
While there is a certain amount of disagreement at the specific and subspecific levels,
the four genera appear to be rather well defined, even if their relationships with one another
are generally none too clear. Although the Helmeted Guineafowl has been known to hybridize
with the Vulturine Guineafowl (Acryllium vulturinum) in captivity, the resultant young are
sterile.
Morphological Aspects
Although the guineafowl constitute a distinctive group within the Galliformes, in structure
and morphology they are generally rather similar to most other members of the order.
In the guineafowl, size appears to be related to the density of the vegetation in the
habitats typically occupied by each species. Thus, the White-breasted, Black and Plumed
Guineafowl, the species that frequent the areas with the thickest vegetation cover, are the
smallest, with average wing lengths of about 20.5 cm for the first two species, and 22.5 cm in
the last. The Crested Guineafowl, which inhabits areas with medium to high vegetation
density, has a mean wing length of 24.5-26 cm, whereas the equivalent figure for the
Helmeted Guineafowl of open or relatively open habitats is 25-28 cm.
Finally, the largest species, the Vulturine Guineafowl, occupies open, arid ground and has a
mean wing length of 29.5 cm.
Like other Galliformes, guineafowl are rather corpulent birds. The oval body is
voluminous, exaggerating the bird’s relatively small head. In all species, both the head and the
neck are virtually featherless, and the bare skin is often beautifully coloured. The bare skin
appears to play an important role in thermoregulation. It is also contorted into a whole series
of adornments, including cartilaginous gape wattles, notably in the Plumed Guineafowl, folds
of skin, for instance on the ind-neck of the Crested Guineafowl, warts on the cere, and
cartilaginous bristles. As their vernacular names suggest, the Helmeted Guineafowl has a
bony “helmet”, whereas the Plumed and Crested Guineafowl have bushy crests.
The long, incredibly slender neck of the Vulturine Guineafowl has almost certainly
developed in order to enhance the bird’s visibility over vegetation. The aspect is strikingly
exaggerated by a series of long, elegant ornamental plumes which hang down over the breast
and upper back.
All these adornments are amongst the most distinctive features of the different species
or, in some cases, subspecies. The many races of the Helmeted Guineafowl are generally
separated on the length and shape of the bony casque and of the gape wattles, and also on the
presence or absence of cere bristles and filoplumes on the hindneck.
The bill is usually short and stout, with the upper mandible curved. It is particularly
feeble in the Black Guineafowl, but, in contrast, appears really quite robust in the Vulturine
Guineafowl, an effect again enhanced by the very thin neck.
The wings are rather small and rounded, clearly belonging to non-migratory birds.
They are designed for short bursts of rapid flight, enabling the birds to make a quick escape
when danger threatens, but are quite unsuitable for any form of sustained flight. This is again
typical of galliform birds, as is their style of flight, involving an explosive take-off and a
series of rapid, powerful wingbeats, followed by a glide down into cover. In line with the
limited flying abilities of the guineafowl is their short tail, composed of 14-16 rectrices. The
exception is the Vulturine Guineafowl, which has a relatively long tail, particularly the central
tail feathers.
The tarsus is strong and generally fairly short, although once again the Vulturine
Guineafowl is an exception, as it has rather long tarsi. The legs and feet, with three front toes,
a single hind one and strong claws, are perfectly adapted to a terrestrial lifestyle, and reflects
the fact that guineafowl mostly move about by walking.
Agelastes is the only genus in which birds sport spurs; these sprout from the
tasometatarsus as in the junglefowl(Gallus), not from the hypotarsal ridge as in most other
Phasianidae.
Males of Agelastes can have one or two spurs per leg, as also do a few females. The Vulturine
Guineafowl commonly has a strange series of spur-like bumps along the rear face of the
tarsus, the function of which is unknown. They are generally longer in males than in females,
and vary individually both in size and in number, with up to six recorded. Agelastes again
differs from the other guineafowl genera by having the tarsal scales imbricated and in rows, as
in the francolins. The other genera have the tarsi covered with pentagonal scales which are not
lined up in rows.
Plumage is generally black or dark grey, although in all species, except the two
members of the apparently primitive genus Agelastes, this is heavily spotted or vermiculated
with white. There is sometimes a bluish tinge to the plumage, which is typically produced by
the feather spotting being bluish white, rather than pure white, but there are fairly extensive
areas of rich blue on the underparts of the Vulturine Guineafowl.
In all species, sexual dimorphism is limited merely to males being slightly larger than
females, with no evident differences in plumage, which is in line with their tendency towards
monogamy and also the participation of the male in the care of chicks. During the breeding
season, there is slight protuberance in the cloaca of males, at any rate in the Helmeted
Guineafowl. However, there is no seasonal variation in plumage in any of the species.
Juvenile plumage tends to be rather similar to that of the adult, but generally drabber,
often with a certain amount of brownish barring. However, the juvenile White-breasted
Guineafowl, in addition to appearing generally duller than the adult, has the plumage pattern
more or less inverted, since it is the belly that is white, not the upper breast or upper back, as
in the adult. Most of the rest of the plumage is brownish black, but recent evidence indicates
that the feathered head and neck almost certainly retain the patterned natal plumage until the
bird moults into full adult plumage, as is the case in other guineafowl. The juvenile Black
Guineafowl likewise shows a white belly.
There is very little information available about moult in the Numididae, but the
Helmeted Guineafowl is known to undergo a complete post-breeding moult, with the flight-
feathers apparently replaced in serially descendant order.
Habitat
The natural distribution of the guineafowl is restricted to sub-Saharan Africa, except for an
isolated population of the Helmeted Guineafowl in north-west Morocco. Between them, the
different species occupy almost all available habitat types, from dense rain forest to arid
steppe and semi-desert. They can be found from sea-level up to considerable altitudes, and the
Helmeted Guineafowl even occurs at 3000m.
The White-breasted, Black and Plumed Guineafowl occupy areas of tropical forest,
essentially in the equatorial belt. They are generally restricted to undisturbed primary forest,
although the Plumed also occurs in very mature secondary growth, while the White-breasted
has been seen in selectively logged forest.
The secretive nature of these species, together with the difficulty of their observation, means
that even basic knowledge on habitat use remains rather limited.
In terms of habitat preference, the Crested Guineafowl can be considered somewhat
intermediate, as it occurs in secondary and riparian forest and also along the forest edge.
Much the most widespread member of the family, the ubiquitous Helmeted Guineafowl, in
addition to its isolated Moroccan population, is found in almost any form of open country
from the Sahel zone southwards. Its most typical habitats include relatively open savanna and
mixed areas of savanna and bush, sometimes interspersed with cultivation, but it is also
common in savanna woodland and along forest edge. Finally, the Vulturine Guineafowl is the
species that occurs in the driest, most open habitats. It is typically found in rather arid zones,
including mixed Acacia and Commiphora scrub and semi-desert, although it does also occur
in woodland, thickets and even montane forest.
As can be seen, there is a certain amount of overlap between the habitat preferences of
the different species, and in some cases more than one species can occur in a particular areas.
For instance both the Vulturine and White-breasted Guineafowl coincide with the Crested
Guineafowl, although in very different habitats, separated almost by the full breadth of Africa.
Nevertheless, in such cases of overlap, the different species do not tend to associate with one
another, and only rarely do they mix.
The proximity of water-holes, or other sources of drinking water, seems to be a
limiting factor in the distribution of guineafowl, with the exception of the Vulturine
Guineafowl. This species has an exceptionally long caecum constituting over 23% of the
entire length of the intestines, and this may permit it to regulate and limit water loss. Another
requirement, for all species, is the presence of suitable trees for roosting in at night.
General Habits
All guineafowl species have highly terrestrial lifestyles. They fly only rather infrequently,
normally either up onto their roosts at night, or in order to escape from predators, although in
the latter case they often prefer to run off at great speed.
They are gregarious for most of the year, but when the breeding season approaches,
the flocks disperse, so that only solitary individuals or pairs are seen, or the odd small group
perhaps composed of non-breeders. The flocks are governed by a complex social structure,
and research on the Helmeted Guineafowl has shown that, at least in this species, individuals
can remain in the same flock over a period of several years.
Group size, outside the breeding season, varies quite considerably with the species,
from the Black Guineafowl, which normally forms groups of under ten individuals, to the
Helmeted Guieanfowl, which can gather in large flocks of over 200 birds, although this
species usually occurs in much smaller flocks of 15-40 birds. The other species form flocks of
intermediate size. There is no evidence to suggest that flocks have strictly defined territories,
and the ranges of two neighbouring groups can overlap to some extent. Nevertheless, if , for
example, two flocks of White-breasted Guineafowl meet, a fierce fight is likely to ensue. In
contrast to this, several flocks of Helmeted Guineafowl can coincide at drinking sites,
particularly good feeding areas, preferred dusting sites, or roosts, but these flocks will only
defend their own particular territories on rare occasions, for instance where the population is
exceptionally dense.
The various different species of guineafowl have to be constantly alert to the threat
posed by their many potential predators, although there are different ways of dealing with
such threats when they actually materialize. For instance, when a predator approaches a flock
of White-breasted Guineafowl, they separate and scatter until the danger has passed, when
they regroup with the help of a cheeping call. In contrast to this rather passive strategy, a flock
of Helmeted Guineafowl will sometimes perform communal defence against a predator.
The evidence available suggests that all species roost in trees at night, on the whole
communally. Helmeted Guineafowl sometimes use traditional roosting sites over such long
periods that the accumulation of dried out droppings on the ground below has been know to
build up to over 30 cm deep! However, similar persistence has not been recorded for the other
species. Thus, the White-breasted Guineafowl, for example, seems simply to choose a roost-
site every evening in whatever area it happens to be at the end of the day, without there being
the slightest evidence to suggest that such sites may be reused. The Plumed Guineafowl uses a
different tree each night, whereas the Crested Guineafowl may regularly roost in the same
general area. On occasions when there may be no suitable roosting trees in the vicinity, the
Helmeted Guineafowl, at least, will look for some sort of substitute, and this species has even
been observed roosting on telegraph poles.
Despite their apparent reluctance to fly, guineafowl are highly mobile birds, and they
tend to spend the greater part of the day roaming about their territories, searching for food.
For instance, White-breasted Guineafowl spend at least 60% of the day feeding, 20%
preening, 11% in trees or flying, and the rest of their time engaged in other assorted activities.
The daily routine of activities in the Helmeted Guineafowl, the best known species, is
rather well established, varying little from one day to the next. The roosting site is occupied in
the evening, and the birds remain there until dawn, when they set off for their drinking holes,
already starting to feed on the way. Often, they move about in “Indian file”, with all members
of the flock following on behind the dominant male. During the hottest hours of the day, they
give up feeding and concentrate on drinking, and sand- or dust-bathing, which frequently
takes place at traditionally favoured sites. Then they pass on to the business of preening and
oiling. After this they head for a suitable area in the shade of some vegetation, where they
rest, using various devices to reduce body temperature, including gular-fluttering, and also
holding the wings away from the body and moving them from time to time for ventilation. As
the heat of the day begins to ease off, the birds renew their activity and, having revisited the
drinking hole, carry on feeding until it is time to return to the roost. Other activities that are
performed somewhat less regularly include basking in the sun, seeking better shade and day-
time roosting, as well as courtship display and even attempted copulation. In the feeding areas
disputes sometimes break out, and these become increasingly common as the breeding season
approaches.
Voice
Guineafowl are gregarious species that inhabit areas with vegetation that is thick enough to
make visual contact between the different members of a flock rather difficult. They overcome
this problem by means of a full vocal repertoire, which enables a sufficient degree of
communication to meet their ordinary needs. For this reason, most activities have associated
vocalizations, which can vary according to the different forms of behaviour involved.
When a White-breasted Guineafowl comes across a plentiful supply of food, it will
often communicate its discovery to the rest of the flock by means of a call known as “food
muster”, summoning its companions to share the food. The contact calls and leaf scuffing
typically used by this species can be heard about 30 metres away, although some of its other
calls are louder. Apart from feeding and contact calls, other typical call types recorded in
various species include, not surprisingly, those to signal alarm and others used in moments of
aggression.
There are some differences between the sexes in vocalizations. For example, females
of both the Vulturine and Helmeted Guineafowl have a “buck-wheat” call, which they most
typically use during the breeding season, on occasions when they are separated from their
partners.
Breeding
Very little is known about the breeding habits of most members of the Numididae, and in
some cases the size and colour of the eggs constitute virtually the only information available.
Thus, details regarding courtship display, the roles of the sexes, the incubation period, overall
levels of success, and so on, can only be interpolated tentatively from the best know species,
the Helmeted Guineafowl.
Thee seasonality of breeding does not appear to follow particularly well set patterns,
and the genera Agelastes and Guttera can breed at any time of the year. Nevertheless, there do
tend to be certain seasonal peaks which are apparently related to the rains. Thus, the White-
breasted Guineafowl seems to prefer the end of the rains, whereas the Black Guineafowl
probably breeds mostly in the driest months of the year. The other four species generally
show a greater or lesser tendency to nest during the rains. An interesting case is that of the
Crested Guineafowl, as its population living in the vicinity of the equator, where there are two
wet seasons in the year, nest all year round.
The first signs of breeding can be in the form of increased frequency and intensity of
aggressive behaviour, in particular fights between rival males, and the splitting off from the
flock of pairs that have been established. Male Helmeted Guineafowl can associate briefly
with a series of various different females, each in its own turn, over the first four to six weeks.
After this period, the pair stabilizes and then remains firm until the end of the breeding
attempt. As a result of this system, the business of establishing and maintaining the pair-bond
causes considerable expenditure of energy by the male, and this is reflected by a loss of
weight amounting to about 11%. Although there is no clearly defined territory, at any rate in
the case of the Helmeted Guineafowl, throughout this initial period males are perpetually
chasing each other and fighting, and frequent communication, both vocal and visual, between
partners is essential for the maintenance of the pair-bond.
Courtship feeding is known to occur, at any rate in the Helmeted, Crested and
Vulturine Guineafowl, and it is particularly ritualized in the last of these species. All three are
monogamous, although males sometimes attempt to mate with other, unattended females. The
Plumed Guineafowl is probably monogamous too, while individual males of the Black
Guineafowl have been seen with two females.
No nest has ever been found for either the White-breasted or the Black Guineafowl,
but they may well be similar to those of the other members of the family, which are very
simple affairs. Essentially, the nest is a shallow depression in the soil, sometimes at the base
of a tree or in long grass. It may be lined with a few dry leaves, perhaps a little grass, and
sometimes the odd feather.
All species have pale eggs, ranging in colour from dirty white and cream to reddish or
brownish, sometimes with a limited amount of spotting. The eggs are oval, with one end sharp
and the other rounded. The shell is particularly thick, and its surface is pitted, with a dense
covering of pores. It is said that the characteristics of the Guineafowl egg would enable it to
survive undamaged through a grass fire. Clutch size is normally of 4-19 eggs, but even larger
clutches have been recorded, although these probably refer to cases of more than one female
layer in the same nest. There is no record of any species ever laying more than a single clutch
per season.
At any rate in the Helmeted Guineafowl, and probably also in the Crested Guineafowl,
incubation is carried out entirely by the female, and during this period she hardly leaves the
nest at all, while the male stands guard nearby, never letting his partner out of his sight.
Incubation begins once the last egg has been laid, and lasts from 23 days in Guttera and
Acryllium to 24-28 days in umida.
Chicks, or keets, hatch synchronously, a common tendency in birds with nidifugous
chicks, as it reduces the danger of predation. The precocial chicks are led away from the nest
almost immediately, but, although they are capable of feeding themselves, they still require
the guidance and protection of their parents for some time, and the male shares the tasks of
chick-care with the female, at least in the Helmeted Guineafowl. In particular, during the first
12 days of the chicks’ lives, the male takes on the burden of all the day-time brooding, while
the female gradually recovers her strength and the weight she has lost during the formation
and laying of the eggs, and also the entire process of incubation. At night, it is the female’s
turn to take on the entire responsibility of brooding the chicks, and the male goes off to roost
with the rest of the flock. When the chicks are feeding, they are normally accompanied by
both adults, but it is the female that acts as leader of the family group. At 15-20 days old, the
chicks are already able to fly op onto roosts that are two metres off the ground.
When danger threatens, both adults take part in the defence of the chicks, the
Helmeted Guineafowl often performing a frontal display with the wings spread and a
growling threat, before launching into an attack. Guineafowl face a whole host of potential
predators at the various different stages of breeding. For instance, various monkeys, genets
and wild cats sometimes steal eggs, and the last-mentioned can even kill adults at the nest.
Both the Black Crow (Corvus capensis) and the Lanner Falcon (Falco biarmicus) are known
to take Helmeted Guineafowl chicks occasionally.
Once the breeding season draws to a close, the family group is merged into a much
larger flock, consisting of other similar family groups and some non-breeding birds. Within
this larger flock, the unity of the family group tens to persist, and the young birds remain in
the company of their parents for at least two or three months more.
It has been calculated that, in an average population of Helmeted Guineafowl, during a
normal breeding season, about 30% of the birds do not breed, although it is not known
whether or not this percentage refers exclusively to immature birds. These non-breeders hang
about in flocks in the habitual feeding areas, and later join up with the breeders when they
return with their young. Although flocks are fairly stable, each individual does not necessarily
return to the same flock that it belonged to before the breeding season.
Movements
All species of Guineafowl are sedentary, and their movements are limited to the local ones
that they perform daily, over variable circuits, in search of food and water. Although they are
mainly terrestrial, they are surprisingly mobile on their feet, and the area in which they forage
can vary quite considerably from one day to the next, when foraging success is low. Indeed,
apart from the Helmeted Guineafowl, which is known to use traditional roosts, birds may start
and end each day in quite different places.
The long, winding route followed daily by flocks of White-breasted Guineafowl, for
example, takes them, on average, over an area of about 0.9km². The speed at which the birds
move about is directly related to foraging success, varying from 10m/h to 203m/h, with an
average of 115 m/h. In the Helmeted Guineafowl, it appears that larger flocks tend to cover
greater distances daily than smaller ones, although the relative proximity of roosting and
drinking sites are also major factors. In this species, home range has been found to average
about 8.8 km², although it is actually extremely variable depending on the habitat type.
White-breasted Guineafowl
Agelastes meleagrides
French: Pintade à poitrine blanche
German: Weissbrust-Perlhuhn
Spanish: Pintada Pechiblanca
Black Guineafowl
Agelastes niger
French: Pintade noire
German: Schwarzperlhuhn
Spanish: Pintada Negra
Helmeted Guineafowl
umida meleagris
French: Pintade de Numidie
German: Helmperlhuhn
Spanish: Pintada Común
Other common names: West African/Grey-breasted Guineafowl (galeata); Tufted Guineafowl
(mitrata); Reichenow’s Guieanfowl (reichenowi)
Plumed Guineafowl
Guttera plumifera
French: Pintade plumifère
German: Schlichthauben-Perlhuhn
Spanish: Pintada Plumífera
Vulturine Guineafowl
Acryllium vulturinum
French: Pintada Vulturine
German: Geierperlhuhn
Spanish: Pintada Vulturina
Taxonomy. umida vulturina Hardwicke, 1834, West Africa; error = Tsavo, Kenya.
Monotopic.
Distribution. S Ethiopia, NW & SW Somalia and arid parts of N & E Kenya, S to R Pangani
(NE Tanzania)
Descriptive notes. c. 60-72 cm; 1026-1645 g. Larges Guineafowl. Noticeably long neck, legs
and tail; bare skin of head and neck pale bluish grey, with patch of short, dense chestnut down
on nape; well developed black and white hackles; plumage mainly dark grey, with white spots
and vermiculations, and rich blue on underpants; commonly on or more spur-like bumps on
each tarsus. Female similar to male, but averages smaller. Juvenile retains some of streaked
natal down on head; hackles obvious; plumage generally greyish brown, with rufous brown,
buff and black barring and mottling; blue areas of plumage less intensely coloured; lacks
tarsal bumps.
Habitat. Generally found in drier and more open habitats than other Guineafowl, especially
semi-arid Acacia/Commiphora thorn-scrub and grassland with trees and bushes; arid and
semi-arid plateau country. On Mt Marsabit, Kenya, found in montane forest; also in tall
riverine Acacia woodland, when foraging; in areas of sympatry with Guttera pucherani,
presents species sometimes enters dense thickets. Exceptionally ranges as high as 1900 m.
Food and Feeding. Seeds and leaves of grasses and herbs; berries and fruit (Commiphora,
Ficus), roots, bulbs, green buds and shoots. Also takes variety of insects, scorpions, spiders
and small molluscs. Forages on ground, scratching with feet; occasionally climbs up into
bushes and low trees (e.g. Salvadora persica, Commiphora) to feed on berries and fruits.
Unlike other Guineafowl does not appear to require drinking water, and even when water
available in dry season, species does not drink.
Breeding. Probably breeds in or just after rainy season; laying peaks in Jun and Dec-Jan. Nest
is simple scrape on ground, situated in cover of fairly dense grass, rock or bush. Up to 13-15
eggs, laid on successive days; eggs creamy white or pale brown, 49.5-55 x 37.3-41 mm.
Incubation c. 23-25 days; chicks have yellowish buff down with dark brown mottling.
Movements. Sedentary.
Status and Conservation. Not globally threatened. Mace-Lande: