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BULLETIN OF
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ZOOLOGY
VOL. 10
1963
BRITISH MUSEUM (NATURAL HISTORY)
LONDON : 1963
DATES OF PUBLICATION OF THE PARTS
No. i 3° January 1963
No. 2 ...... 14 March 1963
No. 3 29 March 1963
No. 4 ...... 29 March 1963
No. 5 28 May 1963
No. 6 31 May 1963
No. 7 9 July 1963
No. 8 16 July 1963
No. 9 ...... 20 May 1963
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ZOOLOGY VOLUME 10
PAGE
No. I. Ethiopian mites of the genus Androlaelaps Berlese s. lat. (Acari :
Mesostigmata). By W. M. TILL i
No. 2. The Francolins, a study in speciation. By B. P. HALL 105
No. 3. Taxonomy and biology of the genus Lebetus (Teleostei-Gobioidea) .
By P. J. MILLER (PI. i) 205
No. 4. The freshwater gastropod molluscs of Western Aden Protectorate.
By C. A. WRIGHT (Pis. 1-2) 257
No. 5. Observations on the chaetotaxy of the legs in free-living Gamasina
(Acari : Mesostigmata). By G. OWEN EVANS 275
No. 6. A revision of the recent round herrings (Pisces : Dussumieriidae) .
By P. J. P. WHITEHEAD 305
No. 7. The polychaete fauna of South Africa. Part 8 : new species and
records from grab samples and dredgings. By J. H. DAY 381
No. 8. The freshwater gastropod mollusca of Angola. By C. A. WRIGHT
(Pis. 1-16) 447
No. 9. New marine nematodes from off the coast of South Africa. By
WILLIAM G. INGLIS 529
Index Volume 10 553
ETHIOPIAN MITES OF THE
GENUS ANDROLAELAPS
BERLESE s.lat.
(ACARI : MESOSTIGMATA)
W. M. TILL
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 10 No. i
LONDON : 1963
ETHIOPIAN MITES OF THE GENUS
ANDROLAELAPS BERLESE s. lat.
(ACARI : MESOSTIGMATA)
BY
W. M. TILL
British Museum (Natural History)
Pp. 1-104 >' I95 Text-Figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 10 No. i
LONDON: 1963
THE BULLETIN OF THE BRITISH MUSEUM
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ETHIOPIAN MITES OF THE GENUS
ANDROLAELAPS BERLESE s. lat.
(ACARI: MESOSTIGMATA).
By W. M. TILL
British Museum (Natural History], London
CONTENTS
INTRODUCTION
EXTERNAL MORPHOLOGY
CLASSIFICATION
HOST-PARASITE LIST
REFERENCES
INTRODUCTION
The earliest species of the genus Androlaelaps s. lat. (including Haemolaelaps} to be
recorded from the Ethiopian region are those described by Berlese (1911-1918)
and Hirst (1916). Radford (1939-1944) added several more species from this region
and Zumpt & Patterson (1950, 1951) provided the first keys to the Ethiopian species
of the genera Androlaelaps and Haemolaelaps. Since then many new species have
been described by Zumpt and his co-workers, by Keegan (1956) and by Lavoipierre
(1955, 1956). With the increased number of species, the existing keys have been
found to be inadequate as they were based largely on superficial characters which
are subject to intraspecific variation. It was also suspected that many of the
new species might be synonyms of older, imperfectly described species, the types
of which had never been thoroughly re-examined. The aim of the present
investigation is therefore to give redescriptions of the known species, based wherever
possible on type material, to try to evaluate the morphological characters observed
in the different species, and to prepare a key to the females. Several new species
have been included and a host-parasite list is given.
This investigation has been made possible by grants from the British Museum
(Natural History) and from the Wellcome Foundation, London. It is based chiefly
on material in the collections of the British Museum (B.M.N.H.) and the South
African Institute for Medical Research, Johannesburg (S.A.I.M.R.), the latter
collection having been placed at my disposal by Dr. F. Zumpt. Material has also
been received from Dr. P. L. G. Benoit, Musee Royal de 1'Afrique Centrale,
Tervuren, Belgium (M.R.A.C.), Dr. R. Taufflieb, Institut de Recherches Scientifiques
au Congo, Brazzaville (I.R.S.C.) and Miss J. B. Walker, East African Veterinary
Research Organization, Kenya. Types and other specimens have been lent by
Miss M. A. Johnson, Liverpool School of Tropical Medicine, Dr. M. M. J. Lavoipierre,
ZOOL. 10, I I
4 W. M. TILL
George Williams Hooper Foundation, California, Dr. C. D. Radford, Manchester,
and by the Smithsonian Institute, United States National Museum, Washington.
The work was carried out in the British Museum (Natural History) and I am
grateful to the Trustees for providing me with the necessary facilities. My thanks
are due to Dr. G. O. Evans for his guidance and many helpful suggestions, and to
Dr. J. G. Sheals and Mr. D. Macfarlane for advice and assistance in various ways.
EXTERNAL MORPHOLOGY
Females
GNATHOSOMA (figs. 1-5) : The gnathosoma is attached subterminally to the anterior
end of the body. It consists of a cylindrical basis capituli (gnathosomal base)
enclosing the pharynx and chelicerae and bearing antero-laterally a pair of seg-
mented pedipalps. The dorsal wall of the basis capituli extends anteriorly as a
membranous flap known as the tectum. In the genus Androlaelaps this is a trans-
parent structure more or less rounded anteriorly and without ornamentation.
On the ventral surface is a median longitudinal groove, the deutosternum, which
normally bears six rows of anteriorly directed teeth. The number of teeth in each
row varies from one to about six. Anteriorly the ventral wall is produced to form
the hypostome and its processes, the horn-shaped corniculi (external malae) and the
fimbriated hypostomal processes (internal malae). In most Androlaelaps species the
corniculi are prominent and well sclerotized (figs, i & 3), but in A. sangsteri and
A. heliosciuri they are reduced in size, weakly sclerotized and generally incon-
spicuous (fig. 4).
The hypostomal processes show a variety of modifications and are sometimes
very difficult to see. In most Androlaelaps species they are similar to those of
A. murinus (fig. i), the ventral processes having the form of a pair of triangular
flaps, the outer margins of which are fringed. The depth of the fringe varies in
different species and is relatively great in A. tachyoryctes (fig. 2). The hypostomal
processes are unusually large and pilose in A. georychi (fig. 5), whilst in A. walker ae
both the ventral and dorsal processes are modified to form prominent brush-like
structures (fig. 3). In A. heliosciuri the inner margins of the processes are fringed
(fig- 4)-
The ventral surface of the gnathosoma bears four pairs of setae, the anterior
rostral (g.s. i) and the posterior external and internal rostral setae (g.s. 2 and g.s. 3)
on the hypostome, and the capitular setae (g.s. 4) on the basis capituli. The lengths
FIGS. 1-5. Types of gnathosoma (ventral view) in females of the genus Androlaelaps
s. lat.
Fig. i. Androlaelaps murinus (Berlese)
Fig. 2. Androlaelaps tachyoryctes (Radford)
Fig. 3. Androlaelaps walkerae sp nov.
Fig. 4. Androlaelaps heliosciuri sp. nov.
Fig. 5. Androlaelaps georychi sp. nov.
b.c., basis capituli; deut., deutosternum; ext. m., external mala; g.s., gnathosomal
seta; hyp., hypostome; int. m., internal mala; la., labrum.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS RERLESE 5. lat.
6 W. M. TILL
of these setae are relatively constant within the genus. The anterior rostral and
capitular setae are usually subequal in length, except in the scapularis-group, and the
posterior internal rostral seta is 2-3 times as long as the external seta. Lying
dorsal to the hypostome and ventral to the chelicerae is a pointed, ciliated structure,
the labrum.
The chelicerae occupy the dorsal region of the gnathosoma. Each terminates
in a fixed and a movable digit, both of which are normally dentate and well
sclerotized. The fixed digit is usually shorter than the movable one, less strongly
dentate, and bears a transparent, seta-like structure, the pilus dentilis, the form
of which varies in different species. The structure of the chelicerae is peculiar in
the scapularis-group, in that the movable digit is edentate, except for the terminal
hook and at the most one rather blunt subterminal tooth, and bears an additional
cutting process provided with large, outwardly directed teeth. The fixed digit is
reduced, weakly sclerotized, and bears a flagellar pilus dentilis (fig. 170). In the
majority of the Androlaelaps species the pilus dentilis is inflated distal to the point
of insertion (subgenus 1 ' schnolaelaps sensu Fonseca), while in a few, for example
A . hystrici, A . phoenicidi, A . casalis, A . walkerae, it is slightly inflated near its point
of insertion, tapering progressively to a point (subgenus Atricholaelaps sensu
Fonseca) .
On the ventral surface of the chelicera, at the base of the movable digit, is a
semicircle of arthrodial filaments. In most of the Androlaelaps species these are
subequal in length or only slightly variable. In A. heliosciuri (fig. 81) two of
the filaments are considerably longer than the others, and in A. murinus
(fig. 109), and A. zumpti (fig. 195) there is one filament which is strikingly
longer and thicker than the others. The arthrodial membrane between the chelae
appears to form an extension which is usually a small, rounded, transparent flap,
but which, in A . zumpti, is greatly elongated, extending to the tips of the chelae.
The pedipalps are 5-segmented; the first free segment, or trochanter, bears
2 setae, the femur bears 5, the genu 6 and the tibia 14 setae. The tarsus, in addition
to a number of simple setae, bears a two-tined seta-like structure at its base.
IDIOSOMA: The idiosoma is usually ovoid in shape with no sharp demarcation
between the dorsal and ventral surfaces. Almost the entire dorsum is covered by a
single scutum which usually has recticular markings and in some species, e.g. those
of the scapularis-group, may also have a granular appearance. The chaetotaxy of
the dorsal shield follows the basic pattern described by Costa (1961, fig. i) for the
genus Haemolaelaps. Most of the species have the pattern comprising 39 pairs of
setae, although in some species, for example those of the mesopicos-group, one or
more of the marginal pairs may be situated on the adjacent integument and seta
PX$ may be lost. A. longipes has an additional pair of setae (pxi) on the posterior
part of the plate (fig. 93), whilst A. centrocarpus and A. hirsti have extra pairs of
setae (ax and PXI} on both the anterior and posterior parts of the shield. Many
species show some degree of hypertrichy, a variable number of unpaired, accessory
setae developing on the median part of the plate between the "/" series. In
A . later ae and A . villosissimus the dorsal shield has a dense covering of setae which
completely obscures the basic pattern, An intermediate condition occurs in
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE 5. lot. 7
A . walker ae in which the basic pattern is partially obscured by the addition of both
unpaired and paired setae. The setae vary considerably in length and in the
scapularis- and w^so^'cos-groups (excluding A. wilkini] and in A.galagus those on
the central part of the shield are reduced to microsetae.
The ventral surface of the idiosoma bears a sternal, a genital and an anal shield,
all of which may show some degree of intraspecific variation. The sternal shield is
usually broader than long, reticulate in most species but with a granular appearance
in mites of the scapularis-group, and bears two pairs of pores and three pairs of
setae. In most Androlaelaps species the sternal setae are more or less subequal in
length, the second and third pairs being very slightly longer than the first pair. In
A. arvicanthis, A. graingeri, A. marshalli and A. theseus the first pair is relatively
short, being |-| as long as the second pair. Metasternal setae are always present.
The presternal area also has a reticulate pattern and bears the tritosternum, a
bipartite structure of uniform appearance throughout the genus. It consists of a
short, undivided, basal portion and two ciliated arms, the laciniae.
The genital shield consists of a relatively more strongly sclerotized posterior
portion, varying in shape from the slender tapering form seen in A . capensis (fig. 20)
and A. tateronis (fig. 165) to the greatly expanded, flask-shaped form of A.
rhodesiensis (fig. 128), and a more weakly sclerotized anterior portion which forms a
flap covering the genital orifice. The shield usually has a linear or reticulate
pattern at least on the posterior portion. This pattern may be very weakly
developed, as in the scapularis-group, or it may be very clearly defined and extend
on to the anterior portion, as in A. dasymys (fig. 51). In some mites the shield
also has a granular appearance and in all the species it bears only one pair of genital
setae.
The anal shield is ovate or triangular, with the apex directed posteriorly, and
encloses the anal orifice. It bears a pair of paranal setae and a single postanal seta.
The metapodal plates are small sclerotized structures lying posterior to the fourth
pair of coxae. The peritreme extends from the stigma, between coxae III and IV,
to a point over coxa I, and the peritremal plate extends posterior to the stigma for
a distance equal to approximately twice the diameter of the stigma.
The setae arising from the integument of the idiosoma may be simple or barbed
and are variable in number, some species being very sparsely and others relatively
densely haired, as for instance A . taterae and A . georychi. The marginal setae are
defined as those lying between the dorsal shield and an arbitrary line commencing
at the posterior tip of the anal shield and continuing anteriorly on both sides and
equidistant from the lateral margin of the body.
The names of the idiosomal shields have been abbreviated and their measure-
ments taken as follows:—
DL = median length of dorsal shield
DW = greatest width of dorsal shield
SL = median length of sternal shield
SW = width of sternal shield at level of second pair of setae
GWX = greatest width of genital shield
GW2 = width of genital shield at level of genital setae
8 W. M. TILL
AL = median length of anal shield
AW = greatest width of anal shield.
LEGS : The legs consist of six recognizable segments, namely, the coxa, trochanter,
femur, genu, tibia and tarsus, each tarsus terminating in a caruncle with claws.
The segments are usually smooth in outline, but in A. callosus some of them have
prominent, heavily chitinized swellings (fig. 18). Leg II is usually stouter than the
other legs and this feature is most pronounced in the Androlaelaps-group. The
chaetotaxy of the legs follows a basic pattern which is subject to slight variations.
The number of setae per segment found in the majority of the Androlaelaps species,
and which I have taken to be the normal number for this genus, is given in the
following table :—
Leg Coxa Trochanter Femur Genu Tibia Tarsus
I 2 6 13 13 13
II ... 2 5 ii ii 10 16
III ... 2 5 6 9 8 16
IV ... i 5 6 10 10 16
The number of tarsal setae listed does not include the two fine setae at the tip of
the segment.
Variations have been noted in the number of setae on the first and third tibiae
and on the third and fourth genua. Tibia I has 14 setae in A. rhodesiensis and
A. walker ae; tibia III has 9 setae in these two species and also in the species of
the mesopicos-group, in A. centrocarpus , A. hirsti, A. longipes, A. phoeniculi, A. oliffi,
A. tateronis, A. cricetomydis, A. congoensis, A. taterae and A. villosissimus. The last
three species have ten setae on genu III and A. spatuliformis has nine setae on
genu IV. The setae are variable in form and may be smooth or barbed, hair-like,
spine-like, pointed or blunt. In the mites of the Androlaelaps-group one of the
ventral setae on femur II is modified to form a stout spur-like process.
Males
The gnathosoma of the male is similar in structure to that of the female. The
chelicera, however, has its movable digit modified to form a spermadactyl which
serves as a copulatory organ. This may be relatively short and straight, as in
A . casalis (fig. 29) , A . hystrici (fig. 88) and others, or relatively long and coiled as in
A. taterae (fig. 164), A. patersoni (fig. 124) and A. spreo (fig. 145). The fixed digit
bears a pilus dentilis, sometimes very difficult to detect, which resembles the pilus
dentilis of the corresponding female.
The dorsal shield has the same chaetotactic pattern as in the female, although the
setae are often relatively longer.
The ventral shields in most Androlaelaps species are fused to form a single
holoventral plate which shows varying degrees of expansion behind the fourth pair
of legs. The shield bears four pairs of sternal, one pair of genital, and the usual
FIGS. 6-9. Androlaelaps casalis (Berlese). Venter (Fig. 6) and dorsum (Fig. 7) of
deutonymph; dorsum (Fig. 8) and venter (Fig. 9) of protonymph.
ETHIOPIAN MITES OF THE GENUS A NDROLA ELA PS BERLESE s. lat.
TO W. M. TILL
three anal setae. The number of setae in the preanal region varies between two
or three pairs, in the species with a narrow plate, for example the mesopicos-group,
and many, in those species with an expanded plate, for instance A. murinus and
A . later ae. The usual number is five pairs.
The anal shield is separated from the sternito-ventral shield in A. centrocarpus ,
A . hirsti and A . longipes (species with 40 or 41 pairs of setae on the dorsal shield)
and also in A, tauffliebi. In all species the genital orifice is presternal.
The chaetotaxy of the legs follows the same pattern as in the corresponding
females although certain setae, for example in the mesopicos-group, may be modified
to form thickened spines or spurs.
Immature Stages
According to Bregetova (1956) Haemolaelaps females are able to lay eggs with
developed larvae or they may give birth to larvae or even to protonymphs. As
far as the Ethiopian species are concerned only proto- and deutonymphs have been
recorded up to now.
The following description of the nymphal stages is based on specimens of A . casalis
from deep poultry litter in England. The immature stages of the other species will
not be described as so very little material is available.
PROTONYMPH (figs. 8-9) : The gnathosoma resembles that of the female except that
the palps have not yet acquired their full complement of setae. The trochanter bears
one ventral seta, and the femur, genu and tibia bear 4, 5 and 12 setae respectively.
The dorsum has a large anterior podonotal shield bearing eleven pairs of setae,
a smaller, posterior, pygidial shield bearing eight pairs of setae, and three pairs of
small intermediate platelets or mesonotal scutellae. The sternal shield, which
bears three pairs of setae and two pairs of pores, does not extend beyond the middle
of coxae IV. The peritremes are short and do not reach beyond coxae III. There
is no genital shield or genital orifice.
The legs have not yet acquired their full number of setae, those present being
listed in the following table:—
Leg Coxa Trochanter Femur Genu Tibia Tarsus
I ... 2 4 10 8 8
II ... 2 4 8 6 7 15
III ... 2 4 5 6 7 15
IV . i 4 4 6 7 15
DEUTONYMPH (figs. 6-7) : The deutonymph is larger than the protonymph and the
palps and legs have the same number of setae as in the adult stages. The podonotal,
mesonotal and pygidial shields have fused to form a single dorsal scutum which
has a lateral incision on either side, behind seta 57. The number of setae on the
shield is the same as in the adult. On the ventral surface the sternal shield extends
to the level of the posterior border of coxa IV. It bears four pairs of setae (the
sternals and metasternals) and three pairs of pores. The peritremes are long,
reaching coxae I. Metapodal plates are present, and also several small areas of
sclerotization on the median part of the venter,
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. n
CLASSIFICATION
It has long been recognized that the genera Androlaelaps and Haemolaelaps are
very closely related to one another as well as to the genus Hypoaspis Canestrini.
Both Androlaelaps and Haemolaelaps were originally described by Berlese as sub-
genera of Laelaps Koch. Subsequently Haemolaelaps (Berlese, 1916) and then also
Androlaelaps (Vitzthum, 1943; Zumpt, 1950; Zumpt & Patterson, 1950) were
placed as subgenera of Hypoaspis, and more recently both have been given generic
status (Strandtmann, 1949; Zumpt & Patterson, 1951; Keegan, 1956; and others).
There is still some confusion, however, regarding the delimitation of the genus
Haemolaelaps. The genera Atricholaelaps Ewing and Ischnolaelaps Fonseca were
synonymized with Haemolaelaps by Strandtmann (1949), a decision which has
been accepted by most other workers on this group. Fonseca (1959), however,
recommended that Haemolaelaps be kept as a monotypic genus for H. marsupialis
Berlese, the remaining species being allocated to the genus Atricholaelaps with its
subgenus Ischnolaelaps. He based these divisions on the form of the pilus dentilis,
a feature which he himself admits is sometimes difficult to interpret. Eubrachylae-
laps Ewing (= Cyclolaelaps Ewing) and Zygolaelaps Tipton were added to the
synonymy by Zumpt and his co-workers (1951, 1958). Cavilaelaps Fonseca was
included in the synonymy by Zumpt & Patterson (1951), but was not listed by
Zumpt & Till (1961). Its status is still doubtful as I have not been able to examine
the type species. The genus Turkiella Zumpt & Till was synonymized with
Androlaelaps by Keegan (1956).
The genera Androlaelaps and Haemolaelaps resemble one another very closely in
their dorsal and ventral sclerotization and chaetotaxy, in their leg chaetotaxy and
in the structure of the chelicerae. In both genera the sternal shield of the female
is broader than long, the pilus dentilis is inflated or long and slender, and the digits
of the male chelicerae are weakly sclerotized and edentate, the fixed digit being
shorter than the movable one. The only differences between Androlaelaps and
Haemolaelaps are that in females of the former genus leg II is relatively more stout
and one of the ventral setae on femur II is thickened to form a spur-like structure.
These features, however, are not clear-cut. There is a gradation both in the thickness
of the second leg and in the nature of its setae from the Haemolaelaps to the
Androlaelaps species. Furthermore, males of the mesopicos-group of Haemolaelaps
species have a stout spur-like seta in the position corresponding to the Androlaelaps
spur, whilst the females have only a simple seta.
The genus Hypoaspis differs from Androlaelaps and Haemolaelaps in that the
length of the sternal shield of the female usually equals or exceeds the breadth ; the
pilus dentilis is short, slender, scarcely visible, and the digits of the male chelicera
are well sclerotized, equal in length, with dentate internal margins. The genus
Gaeolaelaps1 Tragardh, with Laelaps aculeifer Canestrini as type species, appears to
bear the same relationship to Hypoaspis as Androlaelaps does to Haemolaelaps.
It seems possible that these mites may all have arisen from a common free-living
1 The genus Gaeolaelaps was mentioned in Berlese's note book but was never described. Berlese
referred to the genus Geolaelaps in 1924 (Redia 15 : 254), but without designating a type. Tragardh
(1952, Ark, Zool. 4 ser, 2 : 66) states that Berlese (in MS) had designated L, aculeifer as the type.
12 W. M. TILL
Hypoaspis-\ike ancestor, giving rise on the one hand to species with simple setae
on leg II (Hypoaspis} and on the other to species having an enlarged second leg with
thickened setae (Gaeolaelaps} . Modified, elongate, edentate male chelae could
have developed in the mites of both branches which became adapted to a parasitic
mode of life, giving rise to species of the Haemolaelaps and Androlaelaps types
respectively.
In deciding upon the status of these four genera there seemed to be three
possibilities : —
1. To give all four generic status.
2. On the basis of leg setae to combine Haemolaelaps with Hypoaspis and
Gaeolaelaps with Androlaelaps.
3. On the basis of male chelae, female sternal plate and pilus dentilis, to
combine Haemolaelaps with Androlaelaps and Gaeolaelaps with Hypoaspis.
Of these three propositions the third seems to be the most appropriate and
Haemolaelaps has consequently been synonymized with Androlaelaps. In this
connection it is interesting to note that two species described recently (H. travisi
Delfinado 1961 and H. vietnamensis Grochovskaya and Nguyen-Xuan-Hoe 1961)
as having the Androlaelaps type of leg setation have been allocated to the genus
Haemolaelaps by their respective authors.
Bregetova (1956), in her diagnosis of the genus Androlaelaps, describes the
chelicerae as being similar in structure in both sexes, the male having a spermadactyl
added parallel to the movable digit. This is the type of chelicera found in males
of the genus Hypoaspis. However, Berlese's (1887) drawing of A. hermaphrodita,
the type species of Androlaelaps, shows that the male chelae are elongate and
edentate. The Androlaelaps species described from the Ethiopian region are clearly
co-generic with A. hermaphrodita, whereas those listed by Bregetova (.4. sardous
Berlese, A. karawaiewi Berlese and A. pavlovskii Bregetova) are of the Hypoaspis
type.
In the key which I have given to the females of the genus Androlaelaps s. lot.,
couplets 2 to 5 refer to the species bearing the modified spur-like seta (Androlaelaps-
group), whilst the remaining couplets refer to species previously assigned to the
genus Haemolaelaps (Haemolaelaps-group).
On the basis of morphological features, many of the species which I have included
in the Haemolaelaps-group could be arranged in well-defined sub-groups. A.
cryptomius, A. scapularis and A. tauffliebi, three species parasitic on mole-rats,
form a distinct group having the movable cheliceral digit in two parts, microsetae
on the dorsal shield, and " granular " dorsal and ventral idiosomal shields. The
status of these mites may have to be reconsidered and the group regarded either as a
subgenus of Androlaelaps or even as a distinct genus.
A. taterae and A. villosissimus differ from all other Androlaelaps species in their
dorsal chaetotaxy. A. centrocarpus, A. hirsti and A. longipes form a fairly well-
defined group having 40-41 pairs of dorsal setae, nine setae on tibia III and a separate
anal shield in the male.
The mesopicos-group, comprising four species associated with birds, namely,
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 13
A. haydocki, A. mesopicos, A. steyni and A. wilkini, is characterized by the sinuous
posterior setae, the position of the first pair of sternal setae on the presternal area
and by the presence of nine setae on tibia III.
The zulu-glasgowi-group comprises a number of apparently very closely allied
species (A. dasymys s. lat., A. glasgowi, A. pachyptilae, A. patersoni, A. spreo, A.
suncus and A. zulu) whose status is not quite clear, and it is possible that some of
them are no more than subspecies or varieties.
A. glasgowi represents a cosmopolitan species complex which has been recorded
from a wide variety of mammalian and bird hosts (Strandtmann, 1949 ; Bregetova,
1956). According to Bregetova, this species has only 38 pairs of setae on the dorsal
shield, setae 53 being absent. This has been confirmed in specimens from Japan
and Astrakhan in the collection of the British Museum (Natural History) . American
specimens which I have been able to examine, namely, the type specimen of A.
glasgowi (lent by the Smithsonian Institute, Washington) and several specimens
from the collection of the South African Institute for Medical Research, have 39 pairs
of setae. Mites from Israel (Costa, 1961) and from Southern Africa also have
39 pairs of dorsal setae as well as a number of asymmetrically arranged setae on the
posterior part of the dorsal shield. There are also slight morphological differences,
chiefly in relative measurements, between mites from different host species
(Strandtmann, 1949).
A. zulu very closely resembles A. glasgowi, differing from it in having a larger
genital shield and longer setae on the dorsal shield. In both species the pilus
dentilis is strongly inflated proximally, the terminal portion being slender and
recurved. Specimens previously identified by Zumpt & Till (1956) as forms of
A. glasgowi in which the genital shield is bordered by three pairs of setae would
appear to belong to A . zulu.
The remaining members of this group differ from A. glasgowi and A. zulu, and
from one another, in the form of the pilus dentilis, the relative lengths of the
arthrodial filaments at the base of the movable digit of the chelicera and the dorsal
setae, and the relative measurements of parts of the body.
A. dasymys s. lat. is considered to include A. davisi and A. labuschagnei which
appear to differ from A. dasymys s. sir. only in size, degree of sclerotization and
relative setal lengths. A. dasymys was based on a single damaged specimen which
lacks most of the dorsal setae and of which the pilus dentilis is not clearly visible.
By courtesy of Dr. P. L. G. Benoit (Musee Royal de 1'Afrique Centrale) three mites
from Rhabdomys pumilio have been examined which appear to be identical with
A. dasymys and which are in rather better condition. These mites are relatively
large and robust, the dorsal shield measuring 1025-1064^ by 700-750^.
The type series of A . davisi consists of rather smaller, less heavily sclerotized
mites, in which the dorsal shield measures 822-888{jL by 516-569^. Examination
of further material from the type host of A . davisi shows the dorsal chaetotaxy of
this species to be variable in that accessory setae may be present or absent. Mites
from different host species show a variation in the relative lengths of the dorsal
setae. The type specimens of A . labuschagnei, which unfortunately are not in very
good condition, appear to be intermediate between A . davisi and A . dasymys s. str.
I4 W. M. TILL
A. dasymys s. lat. has been collected from a variety of hosts, but appears to be
restricted to the Muridae.
The remaining species in the genus Androlaelaps are more difficult to arrange in
groups. Whatever feature is used for the purpose of separation, the resulting
groups are heterogeneous with respect to other features.
Family LAELAPTIDAE
Genus Androlaelaps Berlese
Androlaelaps Berlese, 1903, Zool. Anz. 27 : 14; Zumpt, 1950, Parasitology 40 : 299; Zumpt &
Patterson, /. ent. Soc. S. Afr. 1951, 14 : 69.
Haemolaelaps Berlese, 1910, Redia 6 : 216 & 1916, Redia 12 : 170; Strandtmann, ig^g.J.Parasit.
35 : 325I Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 70; Bregetova, 1956, Acad. Sci.,
U.S.S.R. 61 : 84; Fonseca, 1959, Mem. Inst. ButantanS. Paulo (1957/58) 28 : 46 (syn. nov.).
Atricholaelaps Ewing, 1929, Manual of External Parasites : 186.
Eubrachylaelaps Ewing, 1929, Manual of External Parasites : 186.
1 schnolaelaps Fonseca, 1936, Mem. Inst. Butantan S. Paulo 10 : 19.
Cyclolaelaps Ewing, 1933, Proc. U.S. Nat. Mus. 82 : 5.
Turkiella Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 215.
Zygolaelaps Tipton, 1957, /• Parasit. 43 : 367.
Type species: Laelaps (Iphis) hermaphrodita Berlese, 1887
KEY TO THE FEMALES
1. Femur II with a stout, blunt or thorn-like seta on its ventral surface (Androlaelaps-
group) . . 2
- Femur II with only simple setae on its ventral surface (Haemolaelaps-group) . . 6
2. Dorsal shield with seta ax present. Genital shield small, slightly widened behind genital
setae, then tapering posteriorly (figs. 173-174) .... A. theseus Zumpt
- Dorsal shield with seta ax absent. Genital shield large, flask-shaped, broadly rounded
posteriorly ............. 3
3. Dorsal shield with setae px2 and px$ present ....... 4
- Dorsal shield with setae px2 and px$ absent ....... 5
4. Tarsus II terminating in pointed setae only. Genital shield very closely approached to
anal shield. Genital setae relatively short, less than half GW2 (fig. 188).
A. zuluensis Zumpt
Tarsus II with three stout, blunt, terminal spines. Genital shield widely separated from
anal shield. Genital setae relatively long, approximately equal to GW2 (fig. 98).
A. marshalli Berlese
5. Integument of idiosoma with about 21 pairs of relatively short, fine setae (figs. 70-71).
A. graingeri Zumpt & Patterson
- Integument of idiosoma with about 37 pairs of long, coarse setae (fig. 10).
A. arvicanthis Radford
6. Hypostomal processes forming two pairs of prominent brushes, one ventral and one dorsal
(fig. 3) . . . . . . . . . . .A. walkerae sp. nov.
Hypostomal processes fimbriated or pilose, but not forming brushes ... 7
7. Dorsal shield with an overall covering of setae which completely obscures the normal
chaetotactic pattern (fig. 161) . . . . . .A. villosissimus (Berlese)
A. taterae (Zpt. & Patt.)
Dorsal shield with chaetotactic pattern composed of not more than 41 pairs of setae,
excluding the accessory setae on the postero-median part of the shield ... 8
8. Corniculi weakly sclerotized (fig. 4) or reduced, inconspicuous .... 9
- Corniculi well sclerotized, normally developed, conspicuous (figs. 1-3, 5) . . 10
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lai. 15
g. First pair of sternal setae inserted on anterior margin of sternal shield. Distance between
genital and anal shields more than twice distance from anterior margin of anal shield to
anterior margin of anus. Dorsal shield almost parallel-sided (figs. 131-132).
A. sangsteri (Radford)
— First pair of sternal setae inserted on presternal area. Distance between genital and anal
shields about half the distance from anterior margin of anal shield to anterior margin of
anus. Dorsal shield ovoid (figs. 79-80) .... A. heliosciuri sp. nov.
10. Dorsal shield with seta /4 at least 5 times as long as seta J r (fig. 55).
A. galagus (Lavoipierre)
- Dorsal shield with setae /i and /4 subequal . . . . . . . n
11. Setae Z$ whip-like and sinuous (fig. 102). From birds (mesopicos-group) . . 12
- Setae Z$ normal. From mammals and birds . . . . . . . 15
12. Setae 24 and 22 very minute, their lengths about J the distance between their bases, or less.
Sternal shield granular ........... 13
- Setae 14 and 22 longer, their lengths at least £ the distance between their bases. Sternal
shield reticulate (fig. 182-183) ....... A. wilkini (Till)
13. Anterior seta of coxa III thick, spine-like (fig. 148) ... A. steyni (Till)
- Anterior seta of coxa III normal, hair-like . . . . . . . . 14
14. Marginal setae of posterior half of dorsal shield simple (fig. 102) A mesopicos (Radford)
Two pairs of marginal setae (82 and 53) of posterior half of dorsal shield whip-like, sinuous
(fig. 78) A. haydocki (Till)
15. Some leg segments with pronounced chitinized swellings (fig. 18) . A. callosus (Berlese)
— Legs without swellings ........... iG
16. Dorsal shield with seta px i developed (fig. 83) ....... 17
— Dorsal shield with seta pxi absent ......... 19
17. Seta ax present (figs. 31, 83) .......... 18
- Seta ax absent (fig. 93) ....... A. longipes (Bregetova)
1 8. Posterior dorsal shield without or with a few accessory setae restricted to the region /3 to
/5 (fig. 83) . .... .A. hirsti (Keegan)
— Posterior dorsal shield with numerous accessory setae extending anterior to seta/i (fig. 31).
A. centrocarpus (Berlese)
19. Tibia III with 9 setae ........... 20
— Tibia III with 8 setae ........... 24
20. Genital shield greatly expanded, greatest width at least 1-6 times width at level of genital
setae .............. 21
- Genital shield not, or only slightly expanded, greatest width not exceeding 1-3 times width
at level of genital setae . . . . . . . . . . .23
21. Genu III with 10 setae ....... A. congoensis sp. nov.
— Genu III with 9 setae ........... 22
22. Lengths of setae 24 and z-z at least twice the distance between their bases (fig>. 129). Tibia 1
with 14 setae ........ A. rhodesiensis (Zpt. & Patt.)
— Lengths of setae 44 and z-z less than half the distance between their bases (fig. 43). Tibia I
with 13 setae ........ A. cricetomydis sp. nov.
23. Tarsus II terminating in pointed setae only. Lengths of setae 24 and 22 approximately J,
or less, the distance between their bases. Dorsal shield with 37 pairs of setae (fig. 126).
A. phoeniculi (Zumpt & Till)
— Tarsus II with at least one of the terminal setae stout and blunt. Lengths of setae ^4 and
Z2. approximately equal to the distance between their bases. Dorsal shield with 39 pairs
of setae (figs. 115, 166) ....... A. tateronis (Radford)
A. oliffl (Zumpt & Patterson)
24. Tarsus II with 3 stout, blunt terminal spines (fig. 157) ..... 25
Tarsus II terminating in pointed setae only ..... 26
25. Sternal shield granular. Caruncle of tarsus IV normal, its length being i£ times the greatest
width of the tarsus . A. tachyoryctes (Radford)
iC W. M. TILL
- Sternal shield faintly reticulated. Caruncle of tarsus IV elongated, its length being at
least twice the greatest width of the tarsus. .... A. capensis (Hirst)
26. Movable digit of chelicera in two parts, one part being dentate; fixed digit of chelicera
weakly sclerotized, membranous in appearance (figs. 135, 170). Setae 24 and 22 very
minute, their lengths not exceeding J the distance between their bases (fig. 134). (scapularis-
group) . 27
- Movable digit of chelicera normal; fixed digit normally developed, well sclerotized.
Setae 14 and z^ relatively longer, their lengths nearly half the distance between their bases,
or longer ............. 29
27. Genital shield short, widely separated from anal shield (fig. 45) A. cryptomius (Radford)
- Genital shield large, flask-shaped, extending almost to anal shield .... 28
28. Width of sternal shield at most 2 \ times its length. Arthrodial filaments at base of movable
digit of chelicera subequal in length (figs. 133, 135) . . A. scapularis (Berlese)
- Width of sternal shield 3 times its length. Four or 5 of the arthrodial filaments at the base
of the movable digit of the chelicera conspicuously longer and stouter than the others
(figs. 168, 170) A. tauffliebi sp. nov.
29. Genu IV with 9 setae A. spatuliformis (Lavoipierre)
• — Genu IV with 10 setae ........... 30
30. Integument of venter, behind coxae IV, relatively densely haired, about 15 pairs, or more,
of truly vental setae (figs. 59, 107) ......... 31
Integument of venter relatively sparsely haired, about 10 pairs, or less, of truly ventral
setae .............. 32
31. Chelicera with arthrodial filaments subequal in length. Pilus dentilis moderately and
more or less evenly inflated, elongate (fig. 61). Hypostomal processes very large and hairy
(fig. 5) . ..... A. georychi sp. nov.
- Chelicera with one very long arthrodial filament. Pilus dentilis strongly inflated basally,
terminal portion slender, curved (fig. 109). Hypostomal processes normal (fig. i).
A. murinus (Berlese)
32. Setae on dorsal shield very long and thick; seta Ji reaches almost to base of 74; lengths
of setae 24 and 22 at least twice the distance between their bases (fig. 63).
A. ghanensis sp. nov.
— Setae on dorsal shield relatively shorter and more slender; seta /i does not reach base of
/3; lengths of setae 24 and 22 less than if the distance between their bases . . 33
33. Pilus dentilis slender (figs. 27, 91) ......... 34
— Pilus dentilis inflated (figs. 187, 195) ......... 35
34. Genital shield widest anterior to first pair of flanking setae, then tapering posteriorly.
Distance between genital and anal shields more than twice distance from anterior margin
of anal shield to anus (fig. 89) A. hystrici (Zumpt & Till)
— Genital shield widest at level of second pair of flanking setae, broadly rounded posteriorly.
Distance between genital and anal shields not exceeding distance from anterior margin of
anal shield to anus (fig. 25) A. casalis (Berlese)
35. Anal shield i£ times longer than broad (fig. 193). . . A. zutnpti sp. nov.
- Anal shield about as long as broad .... zulu-glasgowi complex
a. Parasites of birds ........... b
— Parasites of mammals .......... d
b. Tarsus IV with length to greatest width not greater than 5:1. Seta /i extends to a
point approximately level with base of seta /2 (fig. 118).
A. pachyptilae (Zumpt & Till)
Tarsus IV with length to greatest width about 7:1. Seta /i extends well beyond
base of /2, at least half way to base of /3 . . . . . . . c
c. Pilus dentilis spatulate, terminating in a short, slightly bent, sharp point (fig. 122)
A. patersoni (Zumpt & Till)
Pilus dentilis with a larger proximal and a smaller distal inflation, the latter terminating
in a very fine point (fig. 143) .... A. spreo (Zumpt & Till)
ETHIOPIAN MITES OF THE GENUS ANDBOLAELAPS BERLESE s. la/. 17
d. Chelae 50-65^ long; 2 or 3 of the arthrodial filaments at the base of the movable
digit distinctly longer than the others. Pilus dentilis as in fig. 53 A. dasymys
(Radford)
- Chelae 30-40(0. long; arthrodial filaments at base of movable digit usually subequal in
length e
e. Pilus dentilis with distal portion inflated (fig. 153). Tarsus IV with length to greatest
width not more than 5:1 . . . . . A. suncus sp. nov.
Pilus dentilis with distal portion slender, usually curved (figs. 67, 187). Tarsus IV
with length to greatest width at least 6:1. . . . . . . f
FIGS. lo-ii. Androlaelaps arvicanthis Radford, female. Venter (Fig. 10) ; dorsum (Fig. n).
/. Setae of dorsal shield relatively short, lengths of 24 and Z2 at most slightly exceed the
distance between their bases. Genital shield bordered by two pairs of setae (figs. 65-
66) A. glasgowi (Ewing)
- Setae of dorsal shield relatively long, lengths of 34 and zz at least i£ times the distance
between their bases. Genital shield bordered by three pairs of setae (figs. 185-186).
A. zulu (Berlesc)
Androlaelaps arvicanthis Radford
Androlaelaps arvicanthis Radford, 1944, Parasitology 35 : 162, fig. 4.
Turkiella arvicanthis, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 225.
Androlaelaps equatoriensis Keegan, 1956, /. Egypt, publ. Hlth. Ass. 31 : 230, fig. 15. (synnov.).
ZOOL. IO, I
i8 W. M. TILL
FEMALE (figs. 10-13) : Deutosternum with 6 rows of teeth, 4 to 5 in each row.
Chelae 84(0, long, both digits dentate; pilus dentilis slender, arthrodial filaments at
base of movable digit subequal in length.
Dorsal shield relatively broad, with an overall reticular pattern, well sculptured,
especially anteriorly. DL— 1235-1 264^; DW= 950-1045^; DL/DW= 1-2-1-3.
The shield bears only 37 pairs of setae, px2 and px% being absent. Seta ri distinctly
longer than seta si ; lengths of setae 24 and Z2 about i| times the distance between
their bases; seta Z$ about ij times as long as seta 75.
Sternal shield with a distinct reticular pattern as well as a fine granulation,
anterior and posterior margins very slightly concave. SL= 114-1 24|A; SW=257(x;
SL/SW=o-4-o-5. First pair of sternal setae relatively short, not reaching posterior
margin of sternal shield ; second and third pairs and metasternal setae considerably
longer.
Genital shield flask-shaped, with a reticular and granular pattern. GWX=
257-284^; GW2—2igy.; GW1/GW2=i-2-i-3. Genital setae relatively long,
approximately equal to GW2, and extending to bases of second pair of flanking setae.
Anal shield with linear and granular markings, anterior margin straight or convex,
anus situated at less than its length from the anterior margin. AL=i8o(j,; AW=
200-205^; AL/AW=o-9. Paranal setae level with posterior half of anus, about
4/5 as long as postanal seta.
Integument of idiosoma with about 37 pairs of long ventral and marginal setae,
many of which are barbed. Metapodal plates broadly oval, length to width about
4 =3-
Chaetotaxy of legs normal, anterior and posterior setae on coxa I subequal in
length. Inner ventral seta on tibia II slightly shorter and stouter than outer seta;
inner ventral seta on genu II slightly shorter and stouter than outer seta, but
longer than in A. marshalli. Tarsus II bears three stout, blunt, terminal setae.
Length of tarsus IV about 7 times its width at the base.
MALE (figs. 14-15): Deutosternum as in female; chelicera as illustrated.
Chaetotaxy of dorsal shield as in female. DL= 1064-1092^; DW= 779-884^1;
DL/DW=i-2-i-4. Holoventral shield reticulate, expanded behind coxae IV,
bearing 28-29 setae, 15-16 being situated in the preanal region. Chaetotaxy of
legs as in female, but inner ventral seta on tibia II stouter, resembling the condition
in A. marshalli females.
HOSTS AND LOCALITIES. Arvicanthis niloticus (Desmarest), Bunyoro, Uganda
(types in B.M.N.H.).
Tatera valida (Bocage), Bunyoro, Uganda (S.A.I.M.R.).
Taiera species, Kerio Valley, Kenya and W. Nile district, Uganda (S.A.I.M.R.).
Rattus natalensis (Smith), an Elephant Shrew (single record) and an unidentified
rodent, Kerio Valley, Kenya (S.A.I.M.R.).
Unidentified rodent, Yei, Sudan (paratype of A. equatoriensis Keegan in B.M.N.H.).
Androlaelaps callosus (Berlese)
Hypoaspis (Haemolaelaps) callosits Berlese, 1916, Redia 12 : 171.
Cyclolaelaps lophuromius Radford, 1939, Parasitology 31 : 243, fig. i (syn. nov.}.
ETHIOPIAN MITES OF THE GENUS AN DRO LA EL A PS BERLESE s. la/. 19
Hypoaspis (Haemolaelaps) lophuromius, Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 70.
Haemolaelaps lophuromius, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa, 10 : 238.
Haemolaelaps sudanicus Zumpt & Till, 1954, ^ev- Ecuat. Ent. Parasit. 2 : 213, fig. 3; Kee^an,
1956, /. Egypt, publ. tilth. Ass. 31 : 237, fig. 22 (syn. nov.).
FEMALE (figs. 16-19): Deutosternum with 6 rows of 2 to 4 fine teeth. Chelae
40-46^1 long; pilus dentilis with basal 4/5 inflated, apex sharply recurved according
13
14
15
FIGS. 12-15. Androlaelaps arvicanthis Radford. Chelicera (Fig. 12) and ventral view of
femur, genu and tibia of leg II (Fig. 13) of female. Chelicera (Fig. 14) and venter
(Fig. 15) of male.
to Keegan (1956) but indistinct in the specimens examined; one of the arthrodial
filaments at the base of the movable digit longer than the others.
Dorsal shield reticulate, relatively broad. DL=6oo-620(j.; DW= 445-486^;
DL/DW=i'3-i'4. It bears 39 pairs of setae and one accessory seta at the level of
/4. Anterior and marginal setae longer and stouter than those on central part
of shield and provided with fine barbs. Lengths of setae 14 and Z2 approximately
half the distance between their bases; seta Z$ about i| times as long as seta /5_
W. M. TILL
Sternal shield with a reticular pattern which is most pronounced antero-laterally
and indistinct on the central part of the shield. SL= 83-93^; SW= 134-145^;
SL/SW=o-6-o-7. Sternal setae stout, first pair the same length as the metasternal
setae, second and third pairs slightly longer.
Genital shield granular and also with a reticulate pattern which is well marked
FIGS. 16-19. Androlaelaps callosus (Berlese), female. Venter (Fig. 16); dorsum
(Fig. 17) ; femur, genu, tibia and tarsus of leg IV (Fig. 18) ; chelicera (Fig. 19).
posteriorly but less distinct anteriorly. GWX= 130^1; GW2= 118-127^. Length of
genital setae less than half GW2.
Anal shield approximately as wide as long, the length and width varying between
103^ and ii4(x. Anus situated at approximately its length from the anterior
margin of the shield; paranal setae level with middle of anus, slightly shorter than
postanal seta.
Integument of idiosoma bears 25 pairs of ventral and marginal setae, the latter
being barbed. Metapodal plates ovoid, their length to width about 2:1.
Chaetotaxy of legs normal ; anterior seta on coxa I about 4/5 as long as posterior
seta. Length of tarsus IV 4! times its width at the base. This species is charac-
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lal. zi
terized by the strongly chitinized swellings on several of the leg segments, namely,
femur, tibia and tarsus of leg I, femur II, tarsus III, tibia and tarsus of leg IV.
MALE: Not known.
HOSTS AND LOCALITIES: Lophuromys flavopimctatus Thomas (=Lophuromys zend)
from Kenya (Berlese, 1916; Keegan, 1956), from Kampala, Uganda (syntype of
C. lophuromius in B.M.N.H.), and from Torit, Sudan (type series of H. sudanicus in
S.A.I.M.R.).
Unidentified rodent from Uganda (B.M.N.H.).
Keegan (1956) records this species from Tachyoryctes sp. and a mixed collection of
Otomys and Arvicanthis species in the Rift Valley Province, Kenya.
Androlaelaps capensis (Hirst)
Haemolaelaps capensis Hirst, 1916, /. zool. Res. i : 79, figs, 12-14.
Hypoaspis (Haemolaelaps} capensis, Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 72.
Haemolaelaps capensis, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa, 10 : 246.
Haemolaelaps spinitarsus Berlese, 1918, Redia 13 : 123; Keegan, 1956, Trans. Amer. micr. Soc.
75 : 314, figs. 11-12; Zumpt & Till, 1958, /. ent. Soc. S. Afr. 21 : 267.
Haemolaelaps mystromys Radford i942a, Parasitology 34 : 300, figs. 3-4 (syn. nov.).
FEMALE (figs. 20-22) : Deutosternum with 6 rows of 3-4 small teeth. Chelae
56-64^ long; pilus dentilis slender; arthrodial filaments subequal in length.
Dorsal shield ovoid, reticulate. DL=656-827[z; DW= 394-500^; DL/DW=
1-6-1-7. It bears 39 pairs of setae of moderate length, one or two unpaired
accessory setae sometimes being present. Lengths of 24 and Z2 about 4/5 the distance
between their bases; anterior and marginal setae a little longer; seta Z$ at least
i \ times as long as /5.
Sternal shield reticulate, merging anteriorly with presternal area, pattern weakly
developed posteriorly. SL= 105-133^; SW= 142-150^; SL/SW= 0-7-0-9.
Genital shield apparently smooth, rather slender, and may be slightly widened
behind the genital setae. GW^ 102-128^; GW2= 94-106^; GW1/GW2=i-i-i-3.
Length of genital setae about 2/3 GW2.
Anal shield pear-shaped, 1-3-1-5 times longer than broad. AL= 114-134^;
AW=76-ioo[z. Anus situated at approximately its length from the anterior
margin. Paranal setae level with middle of anus, about 4/5 as long as postanal seta.
Integument of idiosoma bears 23 pairs of ventral and marginal setae, some of the
latter having fine barbs. Metapodal plates elongate, length to width about 4:1.
Chaetotaxy of legs normal. Anterior seta on coxa I about 4/5 as long as posterior
seta. Tarsus II has 3 stout, blunt, terminal spines and 3 stout ventral spines.
Caruncle of tarsus IV relatively long, its length being 2f times the greatest width
of the tarsus. Length of tarsus IV about 8 times its width at the base.
MALE (figs. 23-24): Chelicera as figured, with a slender pilus dentilis. Dorsal
shield reticulate, chaetotaxy as in female. DL=570[j.; DW=323[x; DL/DW=i-8.
Holoventral shield reticulate, with 23 setae. Integument bears about 23 pairs of
marginal and ventral setae, Chaetotaxy of legs as in female, but femur II with a
stout spur,
22
W. M. TILL
FIGS. 20-24. Androlaelaps capensis (Hirst). Venter (Fig. 20), dorsum (Fig. 21) and
chelicera (Fig. 22) of female, Venter (Fig. 23) and chelicera (Fig. 24) of male,
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 23
HOSTS AND LOCALITIES. Cryptomys hottentotus (Lesson), Grahamstown, Cape
Province (type series in B.M.N.H.) and several localities in Natal and the Transvaal
(S.A.I.M.R.).
Cryptomys holosericeus (Wagner), Kruger National Park, Transvaal (S.A.I.M.R.).
Georychus capensis (Pallas), Knysna, Cape Province (S.A.I.M.R.).
Mystromys albicaudatus (Smith), Albany, Cape Province (types of H. mystromys in
B.M.N.H.).
Androlaelaps casalis (Berlese)
Iphis casalis Berlese, 1887, Acari, Myr. Scorp. Ital. Fasc. 38 N.8, figs. 3-5.
Haemolaelaps casalis, Bregetova, 1956, Acad. Sci. U.S.S.R. 61 : 89 & 99, figs. 143-4.
Hypoaspis oculatus Oudemans, 1915, Arch. Naturgesch. (A) 81 : 134.
Haemolaelaps molestus Oudemans, 1929, Ent. Ber. 8 (169) : 13; Willmann, 1939, Abh. Nat.
Ver. Bremen 31 : 173, figs. 5-7.
Atricholaelaps megaventralis Strandtmann, 1947, Proc. ent. Soc. Wash. 49 : 112.
Hypoaspis freemani Hughes, 1948, Mites associated with stored food products: 129, figs. 173-179;
Strandtmann, 1956, /. Kansas ent. Soc. 29 : 138.
Haemolaelaps haemorrhagicus Asanuma, 1952, Misc. Rep. Res. Inst. Nat. Res. No. 25 : 87, fig. i.
(For further references see Strandtmann & Wharton, 1958, Manual of Mesostigmatid Mites:
34)-
FEMALE (figs. 25-27) : Deutosternum with 6 rows each of 5 fine teeth. Chelae
34-40^ long ; pilus dentilis slender ; arthrodial filaments subequal in length.
Dorsal shield oval, reticulate. DL=62o-7iojj.; DW= 376-429^; DL/DW—
1-5-1-7. It bears 39 pairs of symmetrically arranged setae, as well as a variable
number of unpaired accessory setae between the / series. Setae 34 and Z2 are
longer than half the distance between their bases; the longer posterior setae are
barbed; seta Z$ is ii-i|- times as long as /5.
Sternal shield reticulate; SL= 86-96^; SW= 124-140^; SL/SW=o-7-o-8. First
pair of sternal setae about as long as the metasternal setae, but a little coarser;
second and third pairs of sternal setae slightly longer than first pair.
Genital shield expanded and broadly rounded posteriorly, with a well-defined
reticulate pattern which extends at least as far as the anterior flap of the shield.
GW1=i43-i66[jL; GW2= 96-105^; GW1/GW2=i-4-i-6. Genital setae short, less
than half GW2.
Anal shield reticulate, approximately as long as broad. AL=9O-H4fx; AW=
96-ii4fji. Anus situated at about its length from the anterior margin. Paranal
setae inserted on a level between middle and posterior end of anus, and approximately
the same length as the postanal seta.
Integument of idiosoma bears about 21 pairs of ventral and marginal setae.
Metapodal plates elongate, length 4-7 times the width.
Chaetotaxy of legs normal. Posterior seta on coxa I slightly longer than anterior
seta. Three ventral and 3 apical setae on tarsus II are thickened, but do not form
blunt spines. Setae generally rather fine and of moderate length. Length of
tarsus IV about 6 times width at base.
MALE (figs. 28-29) : Chelicerae as figured, the fixed digit bearing a slender pilus
dentilis. Chaetotaxy of dorsal shield as in the female. DL= 466-485^; DW=
W. M. TILL
27
28
29
FIGS. 25-29. Androlaelaps casalis (Berlese). Venter (Fig. 25), dorsum (Fig. 26) and
chelicera (Fig. 27) of female. Venter (Fig. 28) and chelicera (Fig. 29) of male.
ETHIOPIAN MITES OE THE GENUS ANDROLAELAPS BERLESE s. hit. 25
284-322(0.; DL/DW— 1-5-1-6. Holoventral shield reticulate, expanded behind
coxae IV, bearing 23 setae. Chaetotaxy of legs as in female.
HOSTS AND LOCALITIES : Cosmopolitan, from a wide variety of bird and mammalian
hosts, and also found in straw, hay, detritus, etc. Recorded from the Ethiopian
region from Rattus rattus (Linnaeus) and Xerus inauris (Zimmermann), Transvaal,
and from Sciurus carolinensis Gmelin, Cape Province (S.A.I.M.R.).
Androlaelaps centrocarpus (Berlese)
Haemolaelaps centrocarpus Berlese, 1911, Redia 7 1432; Costa, 1961, Bull. Brit. Mus. (nat.
Hist.) Zool. 8 : 27, figs. 39~44-
Haemolaelaps mauritanicus Hirst, 1925, Bull. Com. Etud. Hist. Sci. : 98, fig. 3 (syn. nov.).
FEMALE (figs. 30-32) : Deutosternum with 6 rows of 2-3 teeth. Chelae 6o[o, long;
pilus dentilis slightly inflated, tapering towards the tip ; arthrodial filaments subequal
in length.
Dorsal shield reticulate, widest at level of setae r6, then tapering gradually
posteriorly. DL= 836-911(0.; DW= 485-523(0.; DL/DW= 1-7-1 -8. The shield bears
additional paired setae in the ax and pxi positions, as well as numerous unpaired
setae on the postero-median part and extending anteriorly as far as setae 25. The
setae are long and subequal in length, except for /5, which are shorter. Lengths
of setae 24 and 22 approximately i| times the distance between their bases; seta
Z5 at least i| times as long as seta /5. Some marginal setae have fine barbs.
Sternal shield reticulate. SL= 105-130(0.; SW= 180-190(0.; SL/SW=o-6-O7.
Metasternal setae relatively short, approximately half as long as the sternal setae.
Genital shield granular, with a weakly defined reticular pattern posteriorly;
slightly widened behind the genital setae, then tapering abruptly. GWX= 143-162(0. ;
GW2= 130-143(0.; GW1/GW2=i-i. Genital setae fairly long, about 4/5 GW2,
reaching to base of second pair of flanking setae.
Anal shield pear-shaped, anterior margin straight. AL= 120-145(1; AW=
110-124(0.; AL/AW=i-i-i-3. Anus situated at less than its length from the
anterior margin. Paranal setae level approximately with middle of anus, very
slightly longer than postanal seta.
Integument of idiosoma bears numerous long ventral and marginal setae, many
of the latter being barbed. Metapodal plates elongate, length to width about 6 : i.
Chaetotaxy of legs normal except that tibia III bears 9 setae. Posterior seta of
coxa I thicker than anterior seta, but approximately the same length. Femur I
has 2 short, stout, spine-like setae on its dorsal surface, and trochanter I has a stout,
spine-like dorsal seta. Leg II has a ventral spine-like seta on the trochanter,
femur and genu, and 2 on the tibia. Tarsus II has 3 spine-like ventral setae, and
3 stout, blunt, apical setae. The upper dorsal seta on trochanter IV is considerably
thicker than the other setae on this segment. Length of tarsus IV nearly 10 times
width at base.
MALE (figs. 33-34) : Chelicerae as figured, with a pilus dentilis resembling that of
the female. Dorsal shield as in female. And shield separated from sternito-
ventral shield, the latter strongly reticulate and bearing 7 pairs of setae, 2 pairs
being placed on the area posterior to coxae IV. Chaetotaxy of legs as in female.
26
W. M. TILL
34
33
FIGS. 30-34. Androlaelaps centrocarpus (Berlese). Venter (Fig. 30), dorsum (Fig. 31)
and chelicera (Fig. 32) of female. Venter (Fig. 33) and chelicera (Fig. 34) of male.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE 5. lat. 27
HOSTS AND LOCALITIES. Berlese recorded this species from " Mures " in British
Somaliland. The above description is based on females from Taterillm species,
Kerio Valley, Kenya (S.A.I.M.R.) and on males from Gerbillus gerbillus Olivier,
Israel (received by courtesy of Dr. M. Costa).
FIGS. 35-37. Androlaelaps congoensis sp. nov., female. Venter (Fig. 35); dorsum
(Fig. 36); chelicera (Fig. 37).
NOTE : The specimens described above have been compared with drawings of the
type made by Dr. G. O. Evans. Haemolaelaps mauritanicus Hirst, from Gerbillus
pyramidum Geoffrey, Mauritania, is most probably a synonym of A. centrocarpus.
It is slightly smaller, its setae are relatively finer, and the ventral setation is more
symmetrical than in A. centrocarpus.
28
W. M. TILL
Androlaelaps congoensis sp. nov.
FEMALE (figs. 35-37): Deutosternum with 6 rows of 3-5 very fine teeth and an
ill-defined 7th row with 4 minute teeth. Chelae 48^ long, pilus dentilis small,
slightly inflated, terminating in a sharp point ; arthrodial filaments short, subequal
in length.
38
39
40
FIGS. 38-41. Androlaelaps congoensis sp. nov., venter (Fig. 38) and chelicera (Fig. 39)
of male.
Androlaelaps cricetomydis sp. nov.. venter (Fig. 40) and chelicera (Fig. 41) of male.
Dorsal shield reticulate. 01^=703-722^; DW= 456-504^; DL/DW=i-4~i'6.
The shield bears the usual 39 pairs of setae as well as 11-13 unpaired accessory
setae between the / series. Lengths of setae 24 and Z2 approximately equal to the
distance between their bases ; seta Z5 twice as long as seta 75. Setae all apparently
simple except seta Z$ which has very fine barbs.
Sternal shield reticulate, merging with presternal area, SL (from level of first
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BEKLESE 5. la/. 29
sternal setae) = 105(0,; SW— 114-120(0.; SL/SW^o-g. Sternal and metasternal
setae subequal in length.
Genital shield patterned posteriorly, greatly expanded. GWX= 205-219(0.;
GW2= 1 14-133(0.; GW1/GW2=i-6-l'8. Genital setae relatively short, slightly
more than half GW2.
Anal shield broader than long. AL= 109(0.; AW= 12410.; AL/AW=o-9. Anus
situated at approximately its length from the anterior margin of the shield. Paranal
setae near posterior margin of anus, extending beyond base of postanal seta and
about 3/5 as long as postanal seta.
Integument of idiosoma bears 21 pairs of ventral and marginal setae, all apparently
simple. Metapodal plates elongate, length about 4 times the breadth.
Genu III bears 10 setae and tibia III has 9. Anterior seta on coxa I is about 2/3 as
long as posterior seta. Posterior ventral seta on femur IV extremely long, about
3/5 the total length of the femur. Tarsus IV long and slender, its length about
7 times its width at the base.
MALE (figs. 38-39) : Deutosternum with 4 rows of very fine teeth, the 5th and
6th rows ill defined. Fixed digit of chelicera bears a short pilus dentilis, as in the
female; spermadactyl about Sojo. long.
Dorsal shield 560(0, long, 330(0, wide. Holoventral shield bears 23 setae. Chaeto-
taxy of legs as in female.
HOSTS AND LOCALITIES: Holotype ($), allotype (<^) and 7 female paratypes
(1933.11.14.309-318 part) collected by F. W. ]. Cox from the nest of Spermestes
cucullatus Swainson at Mulungu, Belgian Congo, 5 November, 1919. This species
appears to be very common in birds' nests and has also been found in the nests of
a serin finch, weaver, oriole, sparrow, bunting and wagtail, all from Zomba,
Nyasaland. The specimens are all in the collection of the British Museum (Natural
History) .
Androlaelaps cricetomydis sp. nov.
FEMALE (figs. 42-44) : Deutosternum bears 6 rows of 3-4 small teeth. Chelae
about 45(0. long; pilus dentilis inflated; arthrodial filaments subequal in length.
Dorsal shield reticulate, widest between the posterior setae Si and 82. DL=
850-883(0.; DW= 560-599(0.; DL/DW=i-5. The shield bears 39 pairs of setae,
accessory setae being absent ; setae on central part of shield relatively short, lengths
of 54 and Z2 being less than half the distance between their bases. Marginal setae
longer, coarser and barbed; Si more than twice as long as /i, Z$ about i% times as
long as /5.
Sternal shield reticulate, heavily sclerotized. SL= 128-138(0.; SW= 190-195(0.;
SL/SW=o-7.
Genital shield greatly expanded behind the genital setae, posterior margin straight
or slightly concave. The reticulate pattern extends anterior to the genital setae
where it becomes less well defined. GWX= 238-247(0.; GW2= 128-138^; GWX/GW2
= 1-8-1-9. Genital setae relatively short, less than half GW2.
Anal shield slightly broader than long. AL= 133-144(0.; AW= 152(0.. Anus
3o W. M. TILL
situated at less than its length from the anterior margin. Paranal setae level with
posterior margin of anus, about £ as long as postanal seta.
Integument of idiosoma bears about 23 pairs of marginal and ventral setae. The
3 pairs of ventral setae which flank the genital shield are apparently simple, the
remaining ventral and marginal setae are barbed. Metapodal plates approximately
twice as long as broad.
42 43
FIGS. 42-44. Androlaelaps cricetomydis sp. nov. Venter (Fig. 42), dorsum (Fig. 43)
and chelicera (Fig. 44) of female.
Chaetotaxy of legs normal, except that tibia III bears 9 instead of 8 setae.
Tarsus II bears 3 stout, ventral setae; terminal setae not particularly stout.
Anterior and posterior setae on coxa I approximately equal in length. Length of
tarsus IV about 6|- times its width at the base.
MALE (figs. 40-41) : Deutosternum bears 6 rows of 2-5 teeth ; pilus dentilis inflated
basally. Dorsal shield 650^ long, 437^ wide ; chaetotaxy similar to that of female.
Holoventral shield reticulate, expanded behind coxae IV and bearing 23 setae,
5 pairs lying in the region between coxae IV and the anus. Chaetotaxy of legs
similar to that of the female.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 31
HOST AND LOCALITY: The mites, received from Mr. H. J. Disney, were found in
the food store of Cricetomys gambianus Waterhouse, Malengo Highlands, Tanganyika,
30 August, 1960. Holotype female (1962.6.12.1), allotype male (1962.6.12.2)
and ten female paratypes (1962.6.12.3-7) in the collection of the British Museum
(Natural History).
45
46
FIGS. 45-47. Androlaelaps cryptomius (Radford), female. Venter (Fig. 45); dorsum
(Fig. 46); chelicera (Fig. 47).
Androlaelaps cryptomius (Radford)
Ischnolaelaps cryptomius Radford, 1939, Parasitology 31 : 248, fig. 5.
Hypoaspis (Haemolaelaps) cryptomius, Zumpt & Patterson, 1951, J. ent. Soc. S. Afr. 14 : 71.
Haemolaelaps cryptomius, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 241.
Liponyssus lawrencei Zumpt & Patterson, 1951,7. ent. Soc. S. Afr. 14 : 89, fig. 10; Zumpt &
Till, 1953, S. Afr. J . Med. Sci. 18 : 8 (syn. nov.).
FEMALE (figs. 45-47) : Deutosternum provided with 6 rows of teeth, 2 large and
2 small teeth in the first row, i large tooth in the 2nd to 5th rows, and 2 small teeth
in the sixth row. Anterior rostral seta about i| times as long as capitular seta.
Chelae small, about 24^ long. Fixed digit weakly sclerotized, bearing a long,
slender pilus dentilis; movable digit in two parts, one part edentate except for
terminal hook, the other part weakly sclerotized but with large teeth. Arthrodial
filaments not distinct in the specimens examined.
W. M. TILL
Dorsal shield widest at the level of setae ?5, with a reticulate and granular pattern.
DL= 508-530^; DW= 350-385^; DL/DW=i-4-i-5. The shield bears 39 pairs of
setae. Those on the central part are short and fine, the lengths of setae i<\ and Z2
being less than 1/5 the distance between their bases. The anterior and marginal
setae are longer, and seta Z$ is approximately 2.\ times as long as seta /5-
Sternal shield granular, with some reticulation near the lateral margins ; presternal
area reticulate. SL=7o~75[j.; SW= 130-140^ ; SL/SW=o-5-o-6. First pair of
49
FIGS. 48-50. Androlaelaps cryptomius (Radford), venter of male (Fig. 48).
Androlaelaps dasymys (Radford), venter (Fig. 49) and chelicera (Fig. 50) of male.
sternal setae 4/5 as long as second and third pairs : metasternal setae about half as
long as first pair of setae.
Genital shield relatively short, granular, very slightly widened behind the genital
setae, with a weakly defined reticular pattern. GWX= 96-104^; GW2= 84-88^.
Genital setae relatively long, about 2/3 GW2.
Anal shield approximately as long as broad, length and width varying between
94(x and 104^. Anus situated very close to anterior margin. Paranal setae level
approximately with middle of anus, slightly longer than postanal seta, but not
reaching the base of the latter.
33
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s.lat.
Integument of idiosoma bears about 24 pairs of ventral and marginal setae.
Metapodal plates elongate, their length to width about 8 : i.
Chaetotaxy of legs normal. Posterior seta on coxa I very slightly longer and
thicker than anterior seta. The leg setae are all simple and are not modified to form
thickened or blunt spines. Length of tarsus IV 5-6 times its width at the base.
51
52
FIGS. 51-53. Androlaelaps dasymys (Radford), female (from Mystromys albicaudatus)
Venter (Fig. 51); dorsum (Fig. 52); chelicera (Fig. 53*2). Pilus dentilis of specimens
from Tatera afra (Fig. 536) and Otomys sp. (Fig. 53c).
MALE (fig. 48) : Structure of the chelicerae not very distinct in the only specimen
available, but there appears to be a flagellar pilus dentilis as in the female. Dorsal
shield similar to that of the female, but setae on central part relatively longer.
DL=475pi.; DW = 284^; DL/DW=i-7. Holoventral shield expanded behind
coxae IV and bears 21 setae. The shield has an overall reticulate pattern, and also
a granular appearance especially anterior to coxae IV. Chaetotaxy of legs as in
female.
HOSTS AND LOCALITIES: Georychus capensis (Pallas) from Wynberg, Cape Province
ZOOL. IO, I 3
34 W. M. TILL
(types in Dr. Radford's collection and M.R.A.C.), and from Knysna and Drostdy,
Cape Province (S.A.I.M.R.).
Androlaelaps dasymys (Radford) (s. lat.)
[schnolaelaps dasymys Radford, 1939, Parasitology 31 : 245, fig. 2.
Hypoaspis (Haemolaelaps) dasymys, Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 73.
Haemolaelaps dasymys, Zumpt & Till, 1953, Ann. Inst. Med. trap. Lisboa 10 : 247.
Hypoaspis (Haemolaelaps) labuschagnei Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 85,
fig. 7 (syn. nov.).
Haemolaelaps davisi Zumpt & Till, 1956, Z. Parasitenk 17 : 287, figs. 5-8 (syn. nov.).
FEMALE (figs. 51-53) : Deutosternum with 6 rows of 3-4 teeth, one tooth in each
row being larger than the others in some specimens. Chelae 50-65^ long; pilus
dentilis as figured, its terminal portion appearing either swollen or ribbon-like,
depending on the orientation; two or three of the arthrodial filaments distinctly
longer than the others.
Dorsal shield reticulate, with 39 pairs of setae; unpaired accessory setae may be
present (2-6) or absent. Lengths of setae 24 and Z2 at least 4/5 the distance between
their bases, up to 1-4 times this distance in specimens from Tatera afra, Rattus
rattus, Parotomys species, and some specimens from Otomys species. Marginal
setae coarse, barbed; seta Z$ 1-6-2-5 times as long as seta /5. DL= 822-1 064^;
DW=5i6-750|^; DL/DW=i-4-i-7.
Sternal shield reticulate, sharply demarcated from presternal area. SL=
123-163^.; SW=i54-200[x; SL/SW=o-8-i-o. Metasternal setae a little shorter
than first pair of sternal setae, second and third pairs of sternal setae slightly longer
than first pair.
Genital shield with a distinct reticulate pattern extending almost to the anterior
extremity. GWj= 170-209^; GW2=ii4-i65[x; GW1/GW2=i-3-i-5. Length of
genital seta slightly more than half GW2.
Anal shield reticulate anteriorly and laterally, approximately as long as broad.
AL=i27-200[z; AW=i27-i95[x. Anus situated at a little less than its length from
the anterior margin ; paranal setae level with middle of anus, 4/5 as long as postanal
seta and extending beyond its base.
Integument of idiosoma bears 18-28 pairs of ventral and marginal setae, the
latter being longer, coarser, and provided with fine barbs. Metapodal plates vary
in shape from roughly elliptical to almost round, their length 1-4-2-5 times their
width.
Chaetotaxy of legs normal. Posterior seta on coxa I very slightly longer and
thicker than anterior seta. Tarsus II bears three stout ventral setae but no blunt
spines ; trochanter IV bears a stout seta on its dorsal surface. Length of tarsus IV
about 7 times its width at the base.
MALE (figs. 49-50) : Chelicera as figured ; dorsal shield similar to that of female.
DL=684-788[x; DW 418-420^; DL/DW= 1-6-1-9. Holoventral shield reticulate,
expanded behind coxae IV, bearing 23 setae. Chaetotaxy of legs as in female.
HOSTS AND LOCALITIES: Dasymys incomtus (Sundevall), Kakumiro, Uganda
(syntype of /. dasymys in B.M.N.H.).
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lal.
35
56
57
FIGS. 54-58. Androlaelaps galagus (Lavoipierre). Venter (Fig. 54), dorsum (Fig. 55)
and chelicera (Fig. 56) of female. Venter (Fig. 57) and chelicera (Fig. 58) of male.
36 W. M. TILL
Mystromys albicaudatus (Smith) from Basutoland (type series of H. davisi, the
figured specimens) and from Aliwal North, Cape Province (S.A.I.M.R.).
Rattus rattus (Linnaeus) from the Transvaal (type series of H. labuschagnei in
S.A.I.M.R.).
Rattus natalensis (Smith) from Natal, Cape Province, Transvaal, Orange Free State
(S.A.I.M.R.).
Rattus chrysophilus (De Winton) and Rattus namaquensis (Smith) (Zumpt & Till,
1961; material not re-examined).
Otomys irroratus (Brants) and Otomys species from several localities in South Africa
(S.A.I.M.R.).
Otomys saundersiae Roberts (Zumpt & Till, 1961; material not re-examined).
Otomys sloggetti (Thomas), Basutoland (S.A.I.M.R.).
Parotomys brantsi (Smith), Port Nolloth, Cape Province (S.A.I.M.R.).
Parotomys littledalei Thomas, van Rhynsdorp, Cape Province (S.A.I.M.R.).
Tatera afra (Gray), Basutoland and Transvaal (S.A.I.M.R.).
Rhabdomys pumilio (Sparrman), Ngorongoro, Tanganyika (M.R.A.C.).
Lophuromys sikapusi (Temminck), Cameroons (Taufflieb & Mouchet, 1959).
Androlaelaps galagus (Lavoipierre)
Haemolaelaps galagus Lavoipierre, 1955, Ann. trap. Med. Parasit. 49 : 304, figs. 5-6.
FEMALE (figs. 54-56) : Deutosternum with 6 rows each of 2 small teeth. Chelae
54-56^1 long; pilus dentilis inflated, elongate, bent near tip and terminating in a
fine point; arthrodial filaments subequal in length.
Dorsal shield reticulate, bearing 39 pairs of setae. Those on the central part of
the shield are very short and fine ; they are extremely minute and difficult to detect
in the type specimen examined (by courtesy of Dr. M. M. J. Lavoipierre), but are
more conspicuous in the specimens from Cryptomys sp., on which the drawings are
based. Some of the setae are extremely long and barbed, namely, /4, £3-5 and
54-5; seta /4 is at least 5 times as long as /i. DL= 910-960^; DW= 605-709^;
DL/DW=i-3-i-5.
Sternal shield with a reticulate pattern which is most marked antero-laterally.
SL=i45(j,; SW= 205-2 14^; SL/SW=o-6-o-7- First sternal seta i| times as long
as metasternal seta, second and third sternal setae a little longer than first.
Genital shield sculptured posteriorly, greatly expanded behind the genital setae.
GW1= 247-271 {j.; GW2=i52-i70[z; GW1/GW2= 1-5-17.
Anal shield approximately as wide as long. AL=i43-i62(x; AW= 152-170^.
Anus situated at approximately its length from the anterior margin. Parana!
setae inserted on a level between middle and posterior end of anus, about f as long
as postanal seta.
Integument of idiosoma bears 26 pairs of marginal and ventral setae, many of
which are barbed. Metapodal plates variable in shape, length to width about 2:1.
Chaetotaxy of legs normal ; anterior and posterior setae on coxa I approximately
the same length. Trochanter IV has 2 thick spine-like setae on its anterior surface,
and femur IV has a stout ventral seta and a stout dorsal spine. Antero-dorsal seta
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 37
59
FIGS. 59-61, Androlaelaps
georychi sp. nov., female.
(Fig. 60); chelicera (Fig. 61).
60
Venter (Fig.
59) ; dorsum
on this segment barbed, its length exceeding the width of the femur. Length of
tarsus IV about 6 times its width at the base.
MALE (figs. 57-58) : Chelicera with an inflated pilus dentilis which tapers distally.
Dorsal shield as in female, but setae on central part relatively longer, and 5 accessory
setae present between /3 and //j.. DL=694[j.; DW=456[z; DL/DW=i-5- Holo-
ventral shield reticulate, expanded behind coxae IV, bearing 20-21 setae. Chaeto-
taxy of legs as in female; one ventral seta on femur, genu, tibia and tarsus of leg II
stout and spine-like.
HOSTS AND LOCALITIES: Galago alleni Waterhouse, Barombikang Forest, British
Cameroons (type).
38 W. M. TILL
Cricetomys sp., burrow, Kivu, Belgian Congo (M.R.A.C.).
Cricetomys gambianus Waterhouse, Yaounde, Cameroons (Taufflieb & Mouchet,
1959)-
Androlaelaps georychi sp. nov.
FEMALE (figs. 59-61) : Deutosternum with 6 rows of 3-4 very fine teeth, a larger
median tooth in rows 2 to 4. Hypostomal processes very hairy and large, extending
to anterior margin of palp femur. Chelae about 58 [i long; pilus dentilis elongate
(about 40(ji long), moderately inflated; arthrodial filaments subequal in length.
Dorsal shield reticulate, ovoid in shape, widest at level of seta sj. DL= 964-988^ ;
DW=6i8[x; DL/DW=i-6. The shield bears the usual 39 pairs of setae, which are
more or less subequal in length, and 5-6 accessory setae in the postero-median
region. Lengths of setae 24 and Z2 about i-| times the distance between their
bases; seta ^5 i| times as long as seta 75.
Sternal shield reticulate; SL= 143-152^; SW= 185-200^; SL/SW= 0-7-0-8.
Genital shield with a reticulate pattern which extends over the anterior flap;
greatest width of shield 1-4-1-5 times width at level of genital setae (GW2). Length
of genital setae about 4/5 GW2.
Anal shield approximately as broad as long. AL= 143-1 62 [z; AW =148-162^.
Anus situated at less than its length from the anterior margin of the shield. Paranal
setae near posterior margin of anus, extending to a point level with the posterior
tip of the anal shield, their length about f that of the postanal seta.
Integument of idiosoma bears numerous ventral and marginal setae which all
appear to be barbed, except those adjacent to the genital shield. Metapodal plates
short, ovoid, length to width not greater than 2:1.
Chaetotaxy of legs normal. Length of tarsus IV about 8 times its greatest width.
Caruncle IV elongated, its length more than twice the width of tarsus IV and about
i^ times the length of caruncle III.
HOST AND LOCALITY: Four females from Georychus capensis (Pallas), Knysna,
Cape Province, 24 April, 1954. Holotype and one paratype in the collection of the
S.A.I.M.R., Johannesburg, two paratypes (1962.6.12.8-9) in the collection of the
British Museum (Natural History).
Androlaelaps ghanensis sp. nov.
FEMALE (figs. 62-64) : Deutosternum with 6 rows of 2-4 teeth. Chelae about
62 [x long; pilus dentilis moderately inflated; arthrodial filaments subequal in length.
Dorsal shield reticulate, 817-903^ long, 570-580^ wide at the broadest point,
DL/DW=i-4-i-6. The shield bears 39 pairs of setae and one unpaired accessory
seta at the level of setae 74- All the setae except ii and ri are long and thick.
Lengths of setae 24 and Z2 about twice the distance between their bases; seta Z$
about 1 1 times as long as seta 75 ; seta Ji reaches almost to the base of seta 74-
Sternal shield reticulate, approximately as long as broad; SL=i7i-i8o(x;
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 39
SW=i8o[x. Metasternal setae about 2/3 as long as first pair of sternal setae, second
and third pairs of sternal setae slightly longer.
Genital shield widened near its posterior end and with a distinct pattern extending
to its anterior margin. GWX= 143-152^ ; GW2=io5p.; GW1/GW2=i-4. Genital
setae about as long as first pair of sternal setae, slightly shorter than GW2.
63
Venter (Fig. 62) ; dorsum
62
FIGS. 62—64. Androlaelaps ghanensis sp. nov., female.
(Fig. 63); chelicera (Fig. 64).
Anal shield as long as broad or slightly longer. AL= 157-171^;
AL/AW=i-o-i-i. Anus situated at approximately its length from the anterior
margin of the shield. Paranal setae level with posterior margin of anus, about
3/5 as long as postanal seta.
Integument of idiosoma bears about 27 pairs of ventral and marginal setae; the
marginal and outermost ventral setae are all barbed. Metapodal plates oval in
shape, length to width about 5 : 2.
4o W. M. TILL
Chaetotaxy of legs normal. Trochanter IV bears two very stout spines, one on
the lower ventral and one on the upper dorsal surface. Femur IV bears a stout
sword-like basal spine. Length of tarsus IV about 6 times the width at the base.
HOST AND LOCALITY: Four females from a dormouse (Claviglis spec.) in Ghana,
i January, 1911. Holotype (1962.6.12.10) and three paratypes (1962.6.12.11-13) in
the collection of the British Museum (Natural History).
Androlaelaps glasgowi (Ewing)
Laelaps glasgowi Ewing, 1925, Proc. ent. Soc. Wash. 27 : 6.
Haemolaelaps glasgowi, Strandtmann, 1949, /. Parasit. 35 : 343 ; figs. ; Zumpt & Till, 1956,
Z. Parasitenk. 17 : 282, figs. 1-2.
Laelaps californicus Ewing, 1925, Proc. ent. Soc. Wash. 27 : 5.
Laelaps virginianus Ewing, 1925, Proc. ent. Soc. Wash. 27 : 6.
Hypoaspis microti Oudemans, 1926; Ent. Ber. 7 : 101; Bregetova, 1952, Zool. Zh. 31 : 867.
Haemolaelaps mohrae Oudemans, 1928, Ent. Ber. 7 : 374.
Haemolaelaps cricetophilus Vitzthum, 1930, Zool. Jb. Jena, Abt. Syst. 60 : 417, figs.
Laelaps stegemani Hefley, 1935, /. Kansas ent. Soc. 8 : 22.
1 ' schnolaelaps rhabdomys Radford, 1939, Parasitology 31 : 249, fig. 6 (syn. nov.).
Haemolaelaps scalopi Keegan, 1946, Trans. Amer. Micr. Soc. 65 : 71.
Atricholaelaps sigmodoni Strandtmann, 1946, J. Parasit. 32 : 164.
Atricholaelaps strandtmanni Fox, 1947, Ann. ent. Soc. Amer. 40: 580, figs.
Hypoaspis (Haemolaelaps) eos Zumpt & Patterson, 1951, J. ent. Soc. S. Afr. 14 : 79, fig. 2.
(For further references see Strandtmann & Wharton, 1958, Manual of Mesostigmatid Mites : 37)
FEMALE (figs. 65-67) : Deutosternum with 6 rows of 3-5 teeth. Chelae about
36(0, long; pilus dentilis strongly inflated basally, distal portion slender, curved or
recurved; arthrodial filaments usually subequal in length, slightly variable in
populations from Otomys irroratus.
Dorsal shield reticulate, with a " double " margin. DL= 613-653^; DW=
394-436^; DL/DW= 1-5-1-6. The shield bears 39 pairs of setae and 2-3 unpaired
accessory setae between setae /2 and 74- Lengths of setae 24 and 22 slightly exceed
the distance between their bases; seta Z$ about twice as long as 75-
Sternal shield reticulate and granular. SL— 96-113^; SW= 130-155^; SL/SW—
0-75. Genital shield slightly widened behind genital setae, its reticular pattern
extending over the anterior flap. GW1=i23-i4O[x; GW2=96-io5{Jt; GW1/GW2=
1-2-1-4. Genital setae relatively short, about half GW2.
Anal shield approximately as long as broad. AL= 90-109^; AW= 100-105^.
Anus situated at less than its length from the anterior margin ; paranal setae level
with middle of anus, about 2/3 as long as postanal seta.
Integument of idiosoma bears 22 pairs of ventral and marginal setae, the latter
being a little coarser than the former and provided with fine barbs. Two pairs of
ventral setae flank the genital shield. Metapodal plates 3 or 4 times longer than
broad.
Chaetotaxy of legs normal ; posterior seta on coxa I slightly thicker than anterior
seta, but subequal in length. Length of tarsus IV at least 6 times its greatest
width.
MALE (figs. 68-69) : Chelicera as illustrated, pilus dentilis inflated basally. Dorsal
shield reticulate, chaetotaxy as in female. DL=428[x; DW=257[x; DL/DW=i-7.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE ,. /«/.
65
69
68
FIGS. 65-69. Androlaelaps glasgowi (Ewing). Venter (Fig. 65), dorsum (Fig. 66) and
chelicera (Fig. 67) of female. Venter (Fig. 68^ and chelicera (Fig. 69) of male.
42 W. M. TILL
Holoventral shield reticulate, expanded behind coxae IV and bearing 23 setae.
Integument with about 14 pairs of ventral and marginal setae. Chaetotaxy of legs
as in female.
HOSTS AND LOCALITIES: Rhabdomys pumilio (Sparrman) and Otomys irroratus
(Brants) from several localities in Southern Africa (S.A.I.M.R. ; B.M.N.H; types of
/. rhabdomys in Dr. C. D. Radford's collection).
FIGS. 70-73. Androlaelaps graingeri Zumpt & Patterson, female. Venter (Fig. 70);
dorsum (Fig. 71); chelicera (Fig. 72); femur, genu, tibia of leg II (Fig. 73).
Androlaelaps graingeri Zumpt & Patterson
Androlaelaps graingeri Zumpt & Patterson, 1952, /. ent. Soc. S. Afr. 15 : 162, fig. 2.
Turkiella graingeri, Zumpt & Till, 1953, Ann. Inst. Med. trap. Lisboa 10 : 224.
FEMALE (figs. 70-73): Gnathosoma ventral in position in all the specimens
examined, only the free segments of the palps extending beyond the anterior margin
of the idiosoma. Deutosternum with 6 rows of 3-4 teeth, only one tooth visible
in the 5th row. Chelae 74-76^ long, both digits dentate, movable digit with a
pronounced terminal hook which arches over the fixed digit ; pilus dentilis slender,
arthrodial filaments subequal in length.
Dorsal shield subcircular, with a " double " margin, granular and weakly reticulate.
DL=iooo-io50[x; DW= 825-900^; DL/DW=i-2. Chaetotaxy of dorsal shield
similar to that of A . arvicanthis, i.e. setae px2 and px$ missing. Seta ri considerably
longer than seta ii ; seta Z$ about twice as long as seta /5 ; setae «4-*5> 22, /i-/4
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat.
43
rubbed off in all the specimens examined, but are probably subequal with 23, as
they are in the male. Setae generally finer than in A. arvicanthis and apparently
without barbs.
Sternal shield with an overall granulation and very weak reticular pattern.
SL=8o-96[ji; SW= 197-228^; SL/SW=o-4-o-5. First sternal seta relatively
short, 2nd and 3rd sternal and the metasternal setae considerably longer.
74
75
FIGS. 74-76. Androlaelaps graingeri Zumpt & Patterson, venter of male (Fig. 74).
Androlaelaps haydocki (Till), male. Venter (Fig. 75) ; dorsal view of femur and genu of
leg I (Fig. 76).
Genital shield broadly rounded posteriorly but not greatly expanded behind the
genital setae, granular, with a weak reticular pattern. GWX= 2 15-236^; GW2=
i88-200{x; GW1/GW2=i-i— 1-2. Genital setae relatively long, slightly less than
GW2, reaching base of second pair of flanking setae.
Anal shield reticulate, with a pair of lateral, granular patches; approximately
as long as broad. AL= 162-184^; AW= 158-175^1. Paranal setae near posterior
margin of anus, about 4/5 as long as postanal seta.
44
W. M. TILL
Integument of idiosoma with 21-22 pairs of fine setae. Metapodal plates
subcircular.
Chaetotaxy of legs normal. Anterior and posterior setae of coxa I approximately
equal in length. Ventral setae on genu and tibia II similar to those in A . arvicanthis.
Length of tarsus IV about 6| times the width at the base.
77 78
FIGS. 77-78. Androlaelaps haydocki (Till), female. Venter (Fig. 77); dorsum (Fig. 78).
MALE (fig. 74) : Gnathosoma as in female; spermadactyl about 115^ long. Dorsal
shield with chaetotactic pattern similar to that of female; DL=846[x; DW=646[x;
DL/DW=i-3. Holoventral shield granular with a weak reticular pattern, not
expanded behind 4th pair of coxae, bearing 21 setae, of which 4 pairs lie in the preanal
region. Metapodal plates oval, length about 2| times width. Chaetotaxy of legs
as in female.
HOST AND LOCALITY: Unidentified rodent, Kerio Valley, Kenya (type series,
S.A.I.M.R.).
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 45
Androlaelaps haydocki (Till)
Haemolaelaps haydocki Till, 1959, /. ent. Soc. S. Afr. 22 : 426, figs.
Androlaelaps haydocki is very closely allied to A. mesopicos (Radford) and may
prove to be only a subspecies or variety.
FEMALE (figs. 77-78): The female differs from A. mesopicos in that the dorsal
shield bears 38 pairs of setae, and the marginal setae 57, S2, and 53 are long and
79
80
FIGS. 79-81. Androlaelaps heliosciuri sp. nov., female. Venter (Fig. 79); dorsum
(Fig. 80); chelicera (Fig. 81).
sinuous. The genital shield is slightly more expanded posteriorly (GW1/GW2=
1-3-1-5), and the anal shield is slightly more elongate (AL/AW=i-2-i-4).
MALE (figs 75-76) : The male differs from A . mesopicos in having a long dorsal
seta on genu I.
HOSTS AND LOCALITIES : Thripias namaquus (Lichtenstein) , Luanshya (type series)
and Muliashi Area, Northern Rhodesia; Debeete, Bechuanaland ; Mabelikwa, N.
46 W. M. TILL
Transvaal ; Mapalane, Mozambique (S.A.I.M.R. and B.M.N.H.).
Campethera abingoni (Smith), Luanshya, N. Rhodesia (B.M.N.H.).
Androlaelaps heliosciuri sp. nov.
FEMALE (figs. 79-81) : Deutosternum with 6 rows of 2-3 small teeth; corniculi very
weakly sclerotized. Chelae about 4O(x long, edentate except for terminal hook on
movable digit; pilus dentilis slender; two of the arthrodial filaments considerably
longer and thicker than the others.
Dorsal shield reticulate, widest in posterior half at level of seta 52. DL— 618-
637^; DW=385-4O4(u.; DL/DW=i-6. All 39 pairs of setae are present, but rz,
r\ and r6 are off the shield. Lengths of setae 24 and 22 not greater than half the
distance between their bases; seta Z$ about 3 times as long as seta /5.
Sternal shield granular, with a faint reticular pattern as well. SL=76-89[j.;
SW=i28-i33[A; SL/SW=o-6-o-7. First pair of sternal setae situated off the
plate, slightly shorter than 2nd and 3rd pairs ; metasternal setae as long as 2nd and
3rd pairs of sternal setae.
Genital shield reticulate, the pattern extending over the anterior flap. It is
closely approached to the anal shield and widest at the level of the 2nd pair of
flanking setae. GW1= 142-152^; GW2=9o-95^; GW1/GW2= 1-5-17. Genital
setae relatively short, less than half GW2.
Anal shield broader than long, anterior margin straight. AL=ioi-iiO[ji;
AW— 1 20-123 [j.; AL/ AW— 0-8-0-9. Paranal setae situated between middle and
posterior end of anus, about 2/3 as long as postanal seta.
Integument of idiosoma bears about 34 pairs of ventral and marginal setae,
the former being fine and simple, the latter coarser and barbed. Metapodal plates
long and slender, length to width at least 6 : i.
Chaetotaxy of legs normal. Femur IV bears a stout antero-dorsal seta. Length
of tarsus IV about 4 times the width at the base.
HOST AND LOCALITY: Eight females from Heliosciurus gambianus, (Ogilby),
Amani, Tanganyika, 3 February, 1957. Holotype and three paratypes in the collec-
tion of the South African Institute for Medical Research, four paratypes (1962.6.12.
14-17) in the collection of the British Museum (Natural History).
Androlaelaps hirsti (Keegan)
Haemolaelaps hirsti Keegan, 1956, /. Egypt, publ. Hlth. Ass. 31 : 242, figs. 26-30; Costa, 1961,
Bull. Brit. Mus. (nat. Hist.) Zool. 8 : 14, figs. 17-20.
FEMALE (figs. 82-84) : Deutosternum with 6 rows each of 4 small teeth, except
the last row which has 3 teeth. Chelae about 80 [x long ; pilus dentilis inflated basally,
tapering distally; arthrodial filaments subequal in length.
Dorsal shield ovoid, with a " double " margin, widest about the middle, tapering
posteriorly, granular in appearance with faint reticulations. DL= 1092-1188^;
DW= 703-817^; DL/DW=i-4-i-6. The shield bears 41 pairs of setae, one pair
being present in the ax and one pair in the pxi position. One of the specimens
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 47
examined has an unpaired accessory seta between setae /4 ; the remaining specimens
have no accessory setae. The setae on the central part of the shield are relatively
long, the lengths of 24 and Z2 exceeding the distance between their bases. The
posterior terminal setae Z$ are approximately the same length as setae 14 and at
least 2\ times as long as setae 75.
82
FIGS. 82-84. Androlaelaps hirsti (Keegan), female. Venter (Fig. 82); dorsum (Fig. 83);
chelicera (Fig. 84).
Sternal shield granular with a faint reticulate pattern, presternal area reticulate
and slightly granular. SL= 162-180^; SW= 225-257^; SL/SW=o-6-o-8. Sternal
setae long, first pair reaching posterior margin of shield, third pair reaching middle
of coxae IV ; metasternal seta about half as long as second sternal seta.
Genital shield granular, not expanded posteriorly; GW2= 162-180^. Genital
setae relatively long, reaching base of second pair of flanking setae, their length
nearly equal to GW2.
Anal shield as long as, or slightly longer than, broad. AL= 175-190^; AW=
48 W. M. TILL
i62-i75(x; AL/AW=i-o-i-2. Anus less than its length from the anterior margin
of the shield. Paranal setae inserted on a level between middle and posterior margin
of anus, extending beyond base of postanal seta, and slightly longer than this seta.
Integument of idiosoma with numerous long marginal and ventral setae, many
of the former being barbed. Metapodal plates oval, length about twice breadth.
85
FIGS. 85-88. Androlaelaps hirsti (Keegan), venter (Fig. 85) and chelicera (Fig. 86) of
male.
Androlaelaps hystrici (Zumpt & Till), venter (Fig. 87) and chelicera (Fig. 88) of male.
Tibia III has 9 setae instead of the usual 8. Ventral setae on coxa I nearly equal
in length but posterior seta much thicker. Two short, stout, spur-like setae on
femur I ; one stout ventral seta on genu II, two on tibia II, 3 stout ventral and 3 stout
apical setae on tarsus II, two of the apical ones being relatively blunt. Length of
tarsus IV is 7-8 times its width at the base.
MALE (figs. 85-86) : Fixed digit of chelicera with a long slender pilus dentilis.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat.
49
Deutosternum with 6 rows of 2 or 3 teeth. Dorsal shield as in the female, but the
setae are relatively longer. Anal shield separated from the sternito-ventral shield
which bears 7 pairs of setae, 2 pairs being situated on the part posterior to coxae IV.
Chaetotaxy of legs as in the female.
HOSTS AND LOCALITIES : Unidentified rat from Yubo, Equatoria, Sudan (Paratype
in B.M.N.H.). The above description is based on specimens from Gerbillus
Pyramidum Geoffroy from Israel, received by courtesy of Dr. M. Costa.
r5
' /'/ T\ '
r6
\
FIGS. 89-91. Androlaelaps hystrici (Zumpt & Till), female.
(Fig. 90); chelicera (Fig. 91).
Venter (Fig. 89) ; dorsum
Androlaelaps hystrici (Zumpt & Till)
Haemolaelaps hystrici Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 235, figs. 12-16;
Keegan, 1956, /. Egypt, publ. Hlth. Ass. 31 : 239, fig. 23.
FEMALE (figs. 89-91): Deutosternum with 6 rows each of 3 teeth. Chelae 34^
long ; pilus dentilis slender ; arthrodial filaments subequal in length. Anterior rostral
seta at least i| times as long as capitular seta.
Dorsal shield ovoid, posterior margin almost straight. DL= 600-662^; DW=
ZOOL. IO, I 4
5o W. M. TILL
290-362^; DL/DW =1-8-2-3. Only 36 pairs of setae are present on the shield,
r^-rb being inserted on the adjacent integument. Setae simple, of moderate length ;
lengths of 24 and Z2 about f the distance between their bases ; seta Z$ at least 1 1 times
as long as seta 75.
Sternal shield reticulate. SL= 93-1 14(1; SW= 103-134^; SL/SW=o-8-i-o. Setae
relatively short, first seta reaching base of second.
Genital shield slightly granular, with a few indistinct traces of a reticulate pattern,
very slightly widened behind genital setae, then tapering. GW-,^ 93-1 03^;
GW2= 83-93^. Length of genital setae about half GW2.
Anal shield pear-shaped. AL= 93-134^; AW =93-103^; AL/AW= 1-0-1-3.
Anus situated at approximately its length from the anterior margin. Paranal setae
inserted on a level between middle and posterior end of anus and approximately as
long as the postanal seta.
Integument of idiosoma bears about 22 pairs of ventral and marginal setae, of
which two pairs border the genital shield. Metapodal plates extremely narrow,
length to width about 10 : i.
Chaetotaxy of legs normal. Posterior seta on coxa I slightly longer than
anterior seta. Legs relatively slender; length of tarsus IV 8 times the width at
the base.
MALE (figs. 87-88) : Fixed digit of chelicera with a slender pilus dentilis. Dorsal
shield ovoid, 5oo[x long and 263^ wide ; DL/DW=i-9- Chaetotaxy as in the female,
except that r^-r6 are inserted on the shield. Holoventral shield narrow, bearing
19 setae. Metapodal plates broader than in the female. Chaetotaxy of legs as
in female, but some of the setae relatively stouter, for example, femur II has two
broad, spine-like ventral setae, tibia II has one very broad, spear-like ventral seta,
tarsus II has two broad, spine-like, ventral setae, but the apical setae are slender.
HOSTS AND LOCALITIES: Hystrix cristata Linnaeus from Karamoja district, Uganda
(type series in S.A.I.M.R. and B.M.N.H.) and from Njoro, Rift Valley Province,
Kenya (Keegan, 1956).
Keegan (1956) records this species from Arvicanthis niloticus (Desmarest) and
Rattus rattus (Linnaeus) in Egypt, and also an additional specimen, which should
probably be referred to this species, from Crocidura manni Peters, Kano, Nigeria.
Androlaelaps longipes (Bregetova)
Haemolaelaps longipes Bregetova, 1952, Zoo/. Zh. 31 : 867, figs.; Costa, 1961, Bull. Brit. Mus.
(nat. Hist.) Zoo/. 8 : 21, figs. 28-32.
Haemolaelaps namrui Radford, 1954, Fieldiana, Zoo/. 34 : 310, figs. 56-57 (syn. nov.).
Haemolaelaps aegyptius Keegan, 1956, /. Egypt, publ. Hlth. Ass. 31 : 246, figs. 37-38.
FEMALE (figs. 92-94) : Deutosternum with 6 rows of 3-4 small teeth. Chelae
about 55jj, long; pilus dentilis slightly inflated basally, distal 2/3 slender; arthrodial
filaments subequal in length.
Dorsal shield ovoid, with a reticulate pattern. ~DL=yo^[i ; DW=475[x ; DL/DW=
1-5. The shield bears 40 pairs of setae, one pair being present in the PXI position
on the posterior half of the shield. There is also an unpaired seta near the mid-line
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 51
between setae /3 and /4. The setae on the central part of the shield are relatively
long, the lengths of 14 and Z2 being approximately equal to the distance between
their bases. The anterior and marginal setae are longer, some of the longest setae
having fine barbs. Seta Z5 is more than twice as long as seta 75.
Sternal shield reticulate. SL=g^; SW=i47{ji; SL/SW=o«7. Sternal setae
long, the first pair reaching almost to the posterior margin of the shield, second and
92
FIGS. 92-94. Androlaelaps longipes (Bregetova), female.
(Fig. 93); chelicera (Fig. 94).
93
Venter (Fig. 92) ; dorsum
third pairs slightly longer, the 3rd pair extending well beyond the bases of the
metasternal setae. Metasternal seta less than half as long as 3rd sternal seta.
Genital shield with a reticulate pattern, slightly widened behind genital setae,
then tapering posteriorly. GW^ 125(1,; GW2=i20fi.. Genital setae relatively
long, about 9/10 GW2, extending to base of second pair of flanking setae.
Anal shield slightly longer than wide, anterior margin straight. Length (to base
of postanal seta) is 103(1,, greatest width is 98^. Anus approximately equidistant
between anterior margin of shield and postanal seta. Paranal setae situated between
middle and posterior margin of anus, about the same length as the postanal seta.
52 W. M. TILL
Integument of idiosoma bears about 37 pairs of apparently simple ventral and
marginal setae, of which 2 pairs flank the genital shield. Metapodal plates elongate,
7 times as long as broad.
Chaetotaxy of legs normal except that tibia III has 9 setae. Posterior seta on
coxa I is 1 1 times as long and thick as anterior seta. Stout ventral setae are present
on the femur, genu, tibia and tarsus of leg II, three of the terminal setae on tarsus II
are stout and blunt. The dorsal seta on trochanter IV is much thicker than the
95
97
FIGS. 95-97. Androlaelaps longipes (Bregetova), venter (Fig. 95) and chelicera (Fig. 96)
of male.
Androlaelaps marshalli Berlese, venter of male (Fig. 97).
other setae on this segment. Length of tarsus IV about 8 times the width at the
base.
MALE (figs. 95-96): Fixed digit of chelicera (in A. longipes from the U.S.S.R.)
bears a large, tapering pilus dentilis. Dorsal shield similar to that of female.
Holo ventral shield divided into a sternito- ventral shield and a separate anal shield.
The sternito-ventral shield is moderately expanded behind the 4th pair of coxae
and bears 7-8 pairs of setae, 2 or 3 pairs being situated on the part posterior to
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s.lat. 53
coxae IV. The ventral shield is more reduced in the Russian specimen than in the
type specimen of A. namrui, on which the description and drawings are based.
Integument of idiosoma bears about 31 pairs of setae. Chaetotaxy of legs as in
the female.
HOSTS AND LOCALITIES: Meriones rex Yerbury & Thomas, Ta'izz, Yemen (types
of A. namrui in Dr. C. D. Radford's collection).
98
99
FIGS. 98-100. Androlaelaps marshalli Berlese, female. Venter (Fig. 98); dorsum
(Fig. 99) ; ventral view of femur, genu and tibia of leg II (Fig. 100).
Bregetova (1952) recorded A. longipes from Meriones species in the U.S.S.R. (material
in B.M.N.H.).
Keegan's specimens (A . aegyptius) were collected from a variety of hosts at several
localities in Egypt, and Costa (1961) has found this species chiefly on Meriones
species in Israel.
Androlaelaps marshalli Berlese
Laelaps (Androlaelaps) marshalli Berlese, 1911, Redia 7 : 429.
Androlaelaps marshalli, Keegan, 1956, /. Egypt, publ. Hthl Ass. 31 : 227, fig. 13; Costa, 1961,
Bull. Brit. Mus. (nat. Hist.) Zool. 8 : 45, figs. 75~79-
54 W. M. TILL
Hypoaspis (Androlaelaps) africanus Zumpt, 1950, Parasitology 40 : 30, fig. 2.
Turkiella africanus, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 216.
Hypoaspis (Androlaelaps) africanoides Zumpt & Patterson, 1950, S. Afr. J. med. Sci, 15 : 72,
figs. 2-3.
Androlaelaps oudemansi Radford, 1944, Parasitology 35 : 161, fig. 2 (syn. nov.).
Turkiella oudemansi, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 224.
FEMALE (figs. 98-100) : Deutosternum with 6 rows of teeth, 3 to 6 small, finely
pointed teeth in each row. Chelae similar to those of A. arvicanthis, 70-90^ long;
pilus dentilis slender; arthrodial filaments subequal in length.
Dorsal shield with a " double " margin and an overall reticular pattern. DL=
863-1050(1,; DW= 675-763^; DL/DW= 1-2-1-5. The shield bears 39 pairs of setae
as well as additional unpaired setae between /3 and 74- Many of the setae are
barbed. Lengths of setae 24 and Z2 slightly exceed the distance between their
bases; seta Z$ only slightly longer than 75.
Sternal shield with a well-defined reticular pattern and with concave anterior and
posterior margins. SL= 88-123^; SW=i8o-i93(x; SL/SW= 0-5-0-6. First sternal
seta reaches second pore, second and third sternal and metasternal setae distinctly
longer.
Genital shield broadly rounded posteriorly, with an overall recticular pattern.
GW1=228-250(A; GW2=i5o-i75[z; GW1/GW2=i-4-i-6. Genital setae relatively
long, approximately equal to GW2, and extending almost to the base of the second
pair of flanking setae.
Anal shield with a reticular pattern ; anus at slightly less than its length from the
anterior margin; paranal setae situated near posterior half of anus, only slightly
shorter than postanal seta. AL=i23-i66[i,; AW— 150-175^; AL/AW=o-8-i-o.
Integument of idiosoma bears 30-55 pairs of long ventral and marginal setae,
most of which are barbed. Metapodal plates oval, length about twice the breadth.
Chaetotaxy of legs normal, except that the type of A . oudemansi and some of the
other specimens assigned to this species have 9 setae on tibia III instead of the usual
eight. Anterior and posterior setae on coxa I approximately equal in length.
Thorn-like seta on femur II relatively large; inner ventral seta on genu II short and
spine-like. Inner ventral seta on tibia II a little shorter and considerably stouter
than the outer seta in the types of A. oudemansi and A. africanoides, whereas in
specimens previously identified as A. africanus the differences between these two
setae are less striking, as in A. arvicanthis. Tarsus II bears 3 stout, blunt, terminal
setae. Length of tarsus IV about 8 times its width at the base.
MALE (fig. 97) : Deutosternum as in female ; chelicera similar to that of A . arvi-
canthis. Dorsal shield similar to that of the female. DL= 890-980^; DW=
570-6o8(j,; DL/DW=i-6. Holoventral shield strongly reticulate, expanded behind
fourth pair of coxae, and bears 23-31 setae. The number of setae between coxae IV
and the anus is 10-19, but there may be more on one side than on the other.
Chaetotaxy of legs as in the female.
HOSTS AND LOCALITIES: Recorded in Southern Africa from the following
hosts (Zumpt & Till, 1961) :—
Tatera afra (Gray), Gerbillus paeba Smith, Desmodillus auricularis (Smith), Rattus
paedulcus (Sundevall), Rattus natalensis (Smith), Rattus chrysophilus (De Winton),
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 55
Rattus namaquensis (Smith), Rhabdomys pumilio (Sparrman), Lemniscomys griselda
(Thomas), Saccostomus campestris Peters, Steatomys pratensis Peters, Pedetes capensis
(Forster), Cryptomys hottentotus (Lesson) and Oryctolagus cuniculus (Linnaeus).
Specimens from the following East African hosts have been examined: —
Arvicanthis niloticus (Desmarest), Bunyoro, Uganda (type of A. oudemansi in
B.M.N.H.).
FIGS. 101-103. Androlaelaps mesopicos (Radford), female. Venter (Fig. 101); dorsum
(Fig. 102) ; tip of tarsus IV, ventral view (Fig. 103).
Tatera valida (Bocage), Bunyoro, Uganda (S.A.I.M.R.).
Tatera nigricauda Peters, Lali Hills, Kenya (S.A.I.M.R.).
Tatera species, W. Nile District, Uganda (S.A.I.M.R.).
Taterillus emini (Thomas), W. Nile District, Uganda (S.A.I.M.R.).
Tachyoryctes splendens (Riippell), Muguga, Kenya (S.A.I. M.R.).
Unidentified rodent, Kerio Valley, Kenya (S.A.I.M.R.).
56 W. M. TILL
A. marshalli has also been recorded from an unidentified rodent in the Sudan
and from gerbils in Egypt (Keegan, 1956) and from gerbils in Israel (Costa, 1961).
Two series of mites which have been provisionally assigned to A. marshalli have
been obtained from carnivores, namely, Suricata suricatta (Schreber) from Queens-
town, Cape Province (S.A.I.M.R.), and Ictonyx striatus (Perry) from Muguga, Kenya
(S.A.I.M.R.).
The drawings of A . marshalli are based on paratypes of A . africanoides from Tatera
afra (Gray), S. Rhodesia.
Androlaelaps mesopicos (Kadlord)
Haemolaelaps mesopicos Radford, 1942, Parasitology 34 : 192, figs. 14-19; Till, 1959, /. ent. Soc.
S. Afr. 22 : 423, figs.
Hypoaspis (Haemolaelaps) mesopicos, Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 72.
A series of mites from Mesopicos griseocephalus , Natal, has been assigned to this
species (Till, 1959). Although differing slightly from the types, especially in the
male sex, these mites probably represent no more than a form or variety of A.
mesopicos.
FEMALE (figs. 101-103) : Deutosternum provided with 6 rows of 2-4 small teeth.
Chelae about 40^ long; pilus dentilis small, inflated; arthrodial filaments subequal
in length.
Dorsal shield oval in shape, truncate posteriorly, with a reticulate pattern.
DL= 700-735 [x; DW= 420-438^; DL/DW= 1-6-1-7. Not more than 37 pairs of
setae can be distinguished on the shield, seta 53 being absent and seta r6 arising
from the integument adjacent to the shield. Seta p%3 is present in the type but
absent in the specimens from Natal. The marginal setae are variable in size, the
longest of the S setae being more than 3 times as long as the / setae. Seta Z$ is
sinuous and whip-like and about 20 times as long as seta /5. The setae on the
median part of the shield (i2-i$, J series, z series, Zi-Z4, s2-s6) are very minute,
subequal, the lengths of setae 24 and Z2 being less than a quarter the distance
between their bases.
Sternal shield granular, presternal area weakly reticulate. SL=84-io5(x;
SW= 152-158^; SL/SW=o-6-o-7. First pair of sternal setae slightly shorter than
second and third pairs, and lying on the presternal area. Metasternal setae as
long as the 2nd and 3rd sternal setae.
Genital shield slender, weakly reticulate posteriorly; GWj— 96-144^; GW2=
80-96^; GW!/GW2= 1-1-1-3. Genital setae relatively short, less than half GW2.
Anal shield pear-shaped with a reticular pattern antero-laterally, relatively
shorter and broader in the specimens from Natal. AL= 123-150^; AW= 96-105^;
AL/AW=i-2-i-6. Paranal setae level approximately with middle of anus, not
more than 2/3 as long as postanal seta.
Integument of idiosoma bears about 23 pairs of marginal and ventral setae, which
all appear to be simple, except for the long, sinuous, posterior terminal pair.
Metapodal plates slender, slightly curved.
Tibia HI bears 9 setae and tarsi II, III and IV each bear two stout, blunt,
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 57
terminal spines. The anterior and posterior setae on coxa I are approximately
equal in length and thickness. Length of tarsus IV about 4 times the width at
the base.
MALE (figs. 104-106): Fixed digit of chelicera with a small setiform structure
near its base. Dorsal shield reticulate, and also granular in the specimen from
Natal ; shape and chaetotaxy as in the female, the marginal setae, however, being
104 105
FIGS. 104-106. Androlaelaps mesopicos (Radford), male. Venter of type specimen from
Uganda (Fig. 104); venter (Fig. 105) and chelicera (Fig. 106) of specimen from Natal.
considerably longer. In the type, setae 82 and 5$, and in the Natal specimen 84
also, are long and sinuous. DL=6i8-646[ji; DW=35o-36o[i.; DL/DW=i -6-1-7.
Holoventral shield bears 19 setae. Sternal region similar in shape to sternal shield
of female, with a granular appearance and faint reticular markings ; reticular pattern
more distinct on remaining part of shield. Posterior to coxae IV the shield is narrow
with irregular margins.
Integument of idiosoma bears about 16 pairs of marginal and ventral setae, the
58 W. M. TILL
posterior pair being long and sinuous as in the female. In the specimen from Natal
the longest marginal setae are approximately twice as long as in the type specimen.
Metapodal plates as in the female.
The legs bear the same number of setae as in the female, but there are modifications
in the form of some of the setae. On femur II, the lower medio-ventral seta is
modified to form a thickened spur. Tarsi II, III and IV each bear one stout,
terminal spine, and tarsus II has, in addition, two blunt ventral spurs.
107
FIGS. 107-109. Androlaelaps murinus (Berlese), female. Venter (Fig. 107); dorsum
(Fig. 1 08); chelicera (Fig. 109).
HOSTS AND LOCALITIES: Mesopicos griseocephalus ruwenzori Sharpe, Kabale,
Uganda (types in B.M.N.H.).
Mesopicos g. griseocephalus (Boddaert), Pietermaritzburg, Natal (S.A.I.M.R. and
B.M.N.H.).
Androlaelaps murinus (Berlese)
Laelaps (Haemolaelaps] murinus Berlese, 1911, Redia 7 : 432.
Haemolaelaps murinus, Keegan, 1956, /. Egypt, publ. Hlth. Ass. 31 : 233, figs. 16-17.
Ischnolaelaps arvicanthis Radford, 1939, Parasitology 31 : 246, fig. 3.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat.
59
FEMALE (figs. 107-109) : Deutosternum provided with 6 rows of 3-4 teeth. Chelae
52-54(1 long, movable digit with a pronounced terminal hook which arches over the
fixed digit ; pilus dentilis strongly inflated basally, terminal portion slender, curved ;
one of the arthrodial filaments strikingly longer than the others.
Dorsal shield ovoid; DL= 860-912(1; DW= 595-648(1; DL/DW=i-4-i-5. The
shield bears 39 pairs of setae and 2-3 unpaired accessory setae. Lengths of setae
110
112
FIGS. 110-113. Androlaelaps murinus (Berlese), venter (Fig. no) and chelicera
(Fig. in) of male.
Androlaelaps oliffi (Zumpt & Patterson), venter (Fig. 112) and chelicera (Fig. 113) of
male.
24 and Z2 a little less than the distance between their bases; seta Z$ about twice
as long as seta 75.
Sternal shield strongly reticulate; SL= 120-140(1; SW= 166-184(1; SL/SW=
07-0-8. Metasternal setae shorter than first pair of sternal setae.
Genital shield slightly widened posteriorly and having a reticulate pattern which
extends over the anterior flap. GW^ 184-210(1; GW2= 128-140(1; GWl/GW2=
1-4-1-5. Length of genital setae about half GW2.
Anal shield approximately as long as broad; AL= 145-150(1; AW= 145-16051.
60 W. M. TILL
Anus a little less than its length from the anterior margin; paranal setae inserted
slightly behind middle of anus, a little shorter than the postanal seta.
Integument of idiosoma bears about 60 pairs of setae of which about 15 pairs are
truly ventral in position. The marginal setae tend to be stouter than the ventral
ones and have fine barbs.
Chaetotaxy of legs normal. Anterior seta on coxa I 3/4 as long as posterior seta ;
dorsal seta on trochanter IV stout and spine-like ; two of the dorsal setae on femur IV
spine-like. Length of tarsus IV about 7 times its width at the base.
MALE (figs, iio-iu): The chelicerae have the structure indicated in the figure.
Holoventral shield reticulate, expanded behind coxae IV, bearing 31-32 setae.
Integument of idiosoma bears numerous setae. Chaetotaxy of dorsal shield and
legs as in the female. DL=665[x; DW=437[x; DL/DW— 1-5.
HOSTS AND LOCALITIES: Arvicanthis niloticus (Desmarest) from Uganda (type of
/. arvicanthis in B.M.N.H.) and from Kenya (Keegan, 1956).
Otomys species, Uganda (B.M.N.H.).
Otomys irroratus (Brants) and Rattus natalensis (Smith) from Southern Rhodesia
(S.A.I.M.R.).
Oenomys hypoxanthus (Pucheran) from the Belgian Congo (B.M.N.H.) and from the
Cameroons (I.R.S.C.).
Dasymys incomtus (Sundevall) from Kampala, Uganda (S.A.I.M.R.) and from the
Rift Valley Province, Kenya (Keegan, 1956).
Rhabdomys pumilio (Sparrman), Lophuromys flavopunctatus Thomas and Lemnis-
comys striatus (Linnaeus) from the Rift Valley Province, Kenya (Keegan, 1956).
Androlaelaps oliffi (Zumpt & Patterson)
Hypoaspis (Haemolaelaps) oliffi Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 81, fig. 3.
Haemolaelaps oliffi, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 243.
FEMALE: (figs. 114-116) : Deutosternum with 6 rows of 2-4 teeth. Chela 38-40^
long; pilus dentilis slightly inflated; arthrodial filaments subequal in length.
Dorsal shield ovoid, reticulate. DL= 620-644^; DW=368-4iO(x; DL/DW=
I'5~1'7- The shield bears the usual 39 pairs of setae; lengths of setae 24 and Z2
slightly exceed the distance between their bases; seta Z$ is if times as long as
seta 75.
Sternal shield reticulate; SL=96-io5fx; SW= 130-1 35^; SL/SW=o-7-o-8.
Metasternal setae slightly shorter than first pair of sternal setae.
Genital shield granular and with a reticulate pattern which extends over the
anterior flap. The shield is not expanded behind the genital setae, GWj and GW2
both varying between 127^ and i35[x. Length of genital setae about 2/3 GW2.
Anal shield approximately as long as broad, or slightly longer; AL= 123^1 ;
AW=ii4-n8(jL. Anus situated at about i| times its length from the anterior
margin. Paranal setae level approximately with middle of anus, almost reaching
base of postanal seta. Postanal seta about the same length as the paranal setae,
but slightly thicker.
Integument of idiosoma bears 15 pairs of setae, of which 3 pairs border the
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 61
genital shield; some marginal setae are barbed. Metapodal plates slender, length to
width about 7:1.
Chaetotaxy of legs normal except that tibia III bears 9 setae. Posterior seta
on coxa I nearly twice as long and thick as anterior seta. Anterior seta on coxa II
and both setae on coxa III rather stout. Three of the apical setae on tarsus II
are stout and one of these is blunt. Length of tarsus IV about 5| times the width
at the base.
I.' V! I'./ M
'U
FIGS. 114-116. Androlaelaps oliffi (Zumpt & Patterson), female,
dorsum (Fig. 115); chelicera (Fig. 116).
115
Venter (Fig. 114);
MALE (figs. 112-113): Chelicera as figured; dorsal shield reticulate, chaetotaxy
as in the female ; DL = 428-437/4 ; DW = 276/4 ; DL/DW = 1-6. Holoventral
shield reticulate, bearing 23 setae ; these, excluding the anals, are longer and stouter
than those of the integument. Integument bears about 16 pairs of ventral and
marginal setae ; metapodal plates elongate. Chaetotaxy of legs as in the female.
HOSTS AND LOCALITIES : Widely distributed in Southern Africa, mainly on Tatera
afra (Gray) and Gerbillus paeba Smith, and occasionally found on Saccostomus
campestris Peters and Desmodillus auricularis (Smith) (S.A.I.M.R., B.M.N.H. and
M.R.A.C.). Found also in Tanganyika on Tatera afra (Gray) (B.M.N.H.).
62 W. M. TILL
Androlaelaps pachyptilae (Zumpt & Till)
Haemolaelaps pachyptilae Zumpt & Till, 1956, Z. Parasitenk. 17 : 285, fig. 3.
FEMALE (figs. 117-119): Deutosternum with 6 rows each of 3-4 small teeth.
Chelae 40[x long; pilus dentilis inflated basally, distal half slender and curved;
arthrodial filaments subequal in length.
Dorsal shield ovoid, reticulate, widest between setae rj and 57. DL= 604-691^;
117
FIGS. 117-119. Androlaelaps pachyptilae (Zumpt & Till), female,
dorsum (Fig. 118); chelicera (Fig. 119).
118
Venter (Fig. 117);
DW= 385-446^; DL/DW— 1-5-1-6. The shield bears 39 pairs of setae. Those on
the central part of the shield are relatively short, the lengths of setae 24 and zz
being about half the distance between their bases. The anterior and marginal
setae are considerably longer; seta Z$ is barbed and is about 2\ times as long as
seta /5.
Sternal shield with a reticulate pattern, which is more pronounced antero-
laterally, the central part of the shield having a granular appearance. Presternal
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 63
area strongly reticulate. SL=79-ioo[x; SW= 140-158^; SL/SW=o-5-07. First
sternal seta nearly i| times as long as metasternal seta.
Genital shield short, slightly widened behind genital setae and then tapering
posteriorly; a reticulate pattern is present posteriorly but is not very sharply
defined. GW1=ioo-ii4fx; GW2=g2-g6yi; GWJGWg^ 1-1-1-2. Genital setae the
same length as the ventral setae which flank the shield, about half GW2.
Anal shield slightly wider than long. AL= 105-1 14^; AW= 114-123^; AL/AW=
0-9. Anus situated at less than its length from the anterior margin. Paranal
setae level with middle of anus, not more than 2/3 as long as postanal seta. Length
of postanal seta almost equal to its distance from the anterior margin of the shield.
Integument of idiosoma bears 16 pairs of ventral and marginal setae, two pairs
bordering the genital shield and a third pair lying between the genital and anal
shields. The marginal setae are mostly barbed, the ventral ones simple. Metapodal
plates elongate, length to width about 4:1.
Chaetotaxy of legs normal. Posterior seta on coxa I slightly longer than anterior
seta. Length of tarsus IV about 5 times the width at the base.
MALE: Not known.
HOST AND LOCALITY: Pachyptila desolata (Gmelin) from Heard Island (type
series in S.A.I. M.R.).
Androlaelaps pater soni (Zumpt & Till)
Haemolaelaps patersoni Zumpt & Till, 1956, Z. Parasitenk. 17 : 286, fig. 4.
FEMALE (figs. 120-122) : Deutosternum with 6 rows of 3-6 very fine teeth. Chelae
42-46(0, long; pilus dentilis spatulate, terminating in a short, fine, slightly bent
point ; arthrodial filaments subequal in length.
Dorsal shield bears 39 pairs of setae and a variable number (5-12) of unpaired
accessory setae which may extend anterior to setae 25. Lengths of setae 24 and 22
approximately equal to the distance between their bases; seta Z$ at least twice as
long as seta 75. DL= 683-75 3^; DW =420-481^; DL/DW-i-5-1-6.
Sternal shield reticulate; SL= 100-114^; SW= 130-149^; SL/SW= 0-7-0-8.
Metasternal setae a little shorter than first pair of sternal setae.
Genital shield moderately expanded posteriorly, with a reticulate pattern ex-
tending over the anterior flap. GWX= 135-154^; GW2= 96-114^; GWJGW^—
I'3~1'5- Length of genital setae at least half GW2.
Anal shield with an overall reticulate pattern; AL= 114-131^; AW— 127-131^;
AL/AW=o-9~i-o. Anus situated at approximately its length from the anterior
margin of the shield. Paranal setae level with middle of anus, about 4/5 as long as
postanal seta.
Integument bears about 30 pairs of setae. Outer metapodal plates elongate,
length to width 4-5 : i. Chaetotaxy of legs normal. Anterior and posterior
setae on coxa I subequal in length and thickness. Length of tarsus IV about 7 times
the width at the base.
MALE (figs. 123-124) : The chelicerae have the structure shown in the figure, the
fixed digit bearing a pilus dentilis similar to that of the female.
64
W. M. TILL
124
122
123
FIGS. 120-124. A ndrolaelaps patersoni (Zumpt & Till). Venter (Fig. 120), dorsum (Fig.
121) and chelicera (Fig. 122) of female. Venter (Fig. 123) and chelicera (Fig. 124) of
male.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 65
Chaetotaxy of dorsal shield as in the female, but the setae are relatively longer.
DL= 569-586^; DW=324-359(jt; DL/DW=i«5. Holoventral shield reticulate,
expanded behind coxae IV and bearing 23 setae. The integument bears about
15 pairs of setae. Chaetotaxy of legs as in the female.
HOST AND LOCALITY: Gyps coprotheres (Forster) from Rustenburg, Transvaal
(type and other material, S.A.I.M.R.).
125
126
FIGS. 125-127. Androlaelaps phoeniculi (Zumpt & Till), female. Venter (Fig. 125);
dorsum (Fig. 126); chelicera (Fig. 127).
Androlaelaps phoeniculi (Zumpt & Till)
Haemolaelaps phoeniculi Zumpt & Till, 1954; Rev- Ecuat. Ent. Parasit. 2 : 209, figs. 1-2.
FEMALE (figs. 125-127) : Deutosternum with 6 rows of 1-3 teeth. Chelae 30-32(0,
long; pilus dentilis slender; arthrodial filaments subequal in length.
Dorsal shield reticulate, truncate posteriorly, bearing 37 pairs of setae, two pairs
being absent from the anterior part of the shield. Lengths of setae 24 and Z2 do
not exceed | the distance between their bases ; the setae at the anterior end of the
shield are longer, ii being at least twice as long as 24. Posterior terminal seta Z5
barbed, 5-6 times as long as seta /5. DL=5i7-579[j.; DW= 290-340^; DL/DW=
i -6-1-8. "
ZOOL. IO, I
66 W. M. TILL
Sternal shield similar to that of the mesopicos-group, but relatively broader.
SL=62~72[A; SW= 134-156(0.; SL/SW= 0-4-0-5. Anterior and lateral parts of the
shield granular and reticulate, remaining part of shield apparently smooth ; presternal
area reticulate. First pair of sternal setae and metasternal setae subequal in
length, second and third pairs slightly longer.
Genital shield extends almost to the anal shield and is apparently without any
distinct pattern. It is slightly widened behind the genital setae, with its lateral
margins subparallel and its posterior border almost straight. GW1=ioo-ii4(j.;
GWz=88-g6[L', GW1/GW2=i-o-i-3. Genital setae as long as first pair of sternal
setae, about 1/3 GW2.
Anal shield distinctly longer than broad, anterior margin straight or slightly
convex. AL=93-ii4[jt.; AW=72-83[z; AL/AW= 1-1-1-4. Anus situated at less
than half its length from the anterior margin of the shield. Postanal seta about
1 1 times as long as paranal setae, the latter being level approximately with the middle
of the anus.
Integument of idiosoma bears about 29 pairs of ventral and marginal setae; the
ventral setae are short, fine and simple, the marginal ones coarser and mostly
barbed. Metapodal plates elongate, slightly curved, length to width about 8 : i.
Tibia III with 9 setae ; anterior seta on coxa I at most i| times as long as posterior
seta. The leg setae are simple and there are no stout, blunt spurs. Legs short and
stumpy; length of tarsus IV about 4 times the width at the base.
MALE (not figured) : Length 530^, width 340^. Holoventral shield narrow behind
coxae IV, with a constriction anterior to the anal region. It bears 15 setae, the
first pair lying on the presternal area as in the female. Metapodal plates drop-
shaped.
HOST AND LOCALITIES: Phoeniculus purpureus Miller from Potchefstroom,
Transvaal (type series), from Tsessebe, Bechuanaland, and from Northern Rhodesia
(S.A.I.M.R.).
Androlaelaps rhodesiensis (Zumpt & Patterson)
Hypoaspis (Haemolaelaps} rhodesiensis Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 82,
fig. 4.
Haemolaelaps rhodesiensis, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 243.
FEMALE (figs. 128-130) : Deutosternum provided with 6 rows of 3-5 small teeth.
Chelae 62-70^ long ; pilus dentilis slightly inflated, ribbon-like ; arthrodial filaments
subequal in length.
Dorsal shield broadly oval, reticulate, widest about level of seta Si. DL=
825-912^; DW= 586-700^; DL/DW=i-3-i-4. The shield bears the usual 39 pairs
of setae, as well as two accessory setae at the level of 74. The setae are all very
long, some having fine barbs; lengths of setae 24 and Z2 exceed twice the distance
between their bases; seta /5 is about 4/5 as long as seta Z$.
Sternal shield reticulate; SL=i 14-130^; SW= 170-193^; SL/SW= 0-6-0-7.
Metasternal seta about as long as first sternal seta, second and third sternal setae
slightly longer than the first.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 67
Genital shield with a reticulate pattern extending to the anterior flap, greatly
expanded behind the fourth pair of coxae. GWj= 254-300^; GW2= 130-140^;
GWj/GW2= 2-0-2-2. Genital setae relatively short, their length about 2/3 GW2.
Anal shield triangular, broader than long. AL= 150-160^; AW= 170-184^;
AL/AW= 0-8-0-9. Anus slightly more than its length from the anterior margin.
Paranal setae inserted slightly behind middle of anus, 2/3 to 4/5 as long as the
postanal seta.
128
U9
FIGS. 128-130.
Androlaelaps rhodesiensis (Zumpt & Patterson), female. Venter
(Fig. 128); dorsum (Fig. 129); chelicera (Fig. 130).
Integument of idiosoma bears 15 pairs of long ventral and marginal setae, of which
3 pairs flank the genital shield; most of the setae have fine barbs. Metapodal
plates roughly oval, length about twice the width.
Tibia I has 14 and tibia III has 9 setae. Anterior seta on coxa I about 4/5 as
68
W. M. TILL
long as posterior seta. Most of the leg setae are barbed; setae on some segments
stouter than on others, but there are no blunt spines. Length of tarsus IV about
7 times the width at the base.
MALE: Not known.
HOSTS AND LOCALITIES: Saccostomus campestris Peters from several localities in
Southern Rhodesia (S.A.I. M.R.).
Rattus rattus (Linnaeus) and Rattus morio (Trouessart) from Brazzaville, French
Congo (Zumpt & Till, 1961).
Steatomys pratensis Peters from Southern Rhodesia (one female only, Zumpt &
Till, 1961).
131 132
FIGS. 131-132. Androlaelaps sangsteri (Radford) female. Venter (Fig. 131); dorsum (Fig. 132).
Androlaelaps sangsteri (Radford)
Liponyssus sangsteri Radford, 1942, Parasitology 34 : 190, fig. n; Zumpt & Till, 1953, 5. Afr.
J. Med. Sci. 18 : 7.
Haemolaelaps sangsteri, Strandtmann & Wharton, 1958, Manual of Mesostigmatid Mites : 44.
FEMALE: (figs. 131-132) : Deutosternum provided with 6 rows of teeth, 4 teeth in
the first row, 2 in the second and one in each of the remaining rows. Corniculi
reduced, comparatively small and inconspicuous. Chelicerae missing in the type,
the only available specimen.
Dorsal shield narrow with lateral margins subparallel, granular in appearance,
with faint reticulations. DL=5i8pi; DW=257pi; DL/DW=2-o. The number of
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 69
setae is reduced, only 34 pairs being distinguishable with certainty. Those on the
central part of the shield are short, the lengths of setae 24 and 22 not exceeding half
the distance between their bases. The anterior and marginal setae are longer;
seta Z5 is at least 4 times as long as seta /5 and has fine barbs.
Sternal shield granular, anterior margin slightly convex, posterior margin straight,
SL=96(ji; SW=i32(x; SL/SW=o«7. Sternal setae subequal in length, relatively
short, not reaching bases of following setae. Metasternal setae about as long as
sternal setae.
FIGS. 133-135. Androlaelaps scapularis (Berlese), female. Venter (Fig. 133); dorsum
(Fig. 134); chelicera (Fig. 135).
Genital shield granular, with an indistinct reticulate pattern; widest near its
posterior border, between the first and second pairs of flanking setae ; anterior part
of shield rather sharply pointed. GW1=ioopi; GW2=72[x; GWJGWg— 1-4.
Length of genital setae about half GW2.
Anal shield triangular, approximately as long as broad. AL=82[j.; AW=79jji;
Parana! setae level with middle of anus, about 4/5 as long as postanal seta.
Integument of idiosoma bears 26 pairs of ventral and marginal setae, the longer
ones near the posterior end of the body being barbed. Metapodal plates not
visible.
Legs short and stumpy, their chaetotaxy following the typical Androlaelaps
yo W. M. TILL
pattern ; all the setae simple and hair-like. Length of tarsus IV nearly 4 times the
width at the base.
MALE : Not known.
HOST AND LOCALITY : Protoxerus stangeri (Waterhouse) from Bwamba Valley,
Toro, Uganda (type in B.M.N.H.).
Androlaelaps scapularis (Berlese)
Hypoaspis (Haemolaelaps) scapularis Berlese, 1918, Redia 13 : 124; Keegan, i956a, Trans.
Amer. micr. Soc. 75 : 314, figs. 13-14.
1 'schnolaelaps bathyergus Radford, 1939, Parasitology 31 : 250, figs. 7-8 (syn. nov.).
Haemolaelaps eloffi Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 226, figs. 6-9 (syn. nov.}.
Haemolaelaps natalensis Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 230, figs. 10-11
(syn. nov.}.
FEMALE (figs. 133-135) : Deutosternum provided with 6 rows of 2-4 teeth, the
teeth in rows 5 and 6 being very small and inconspicuous ; anterior rostral seta at
least 1 1 times as long as capitular seta. Chelae very small, 18-2 2 fz long; fixed
digit weakly sclerotized, bearing a slender pilus dentilis ; movable digit in two parts,
one part edentate except for the terminal hook and a small subterminal tooth, the
other part dentate but more weakly sclerotized. Arthrodial filaments subequal in
length.
Dorsal shield broadly oval, granular, with a faint reticulate pattern. DL=445-
570(x; DW=28o-390[x; DL/DW=i-3-i-6. The shield bears 39 pairs of setae.
The centrally situated ones are very small, the lengths of 24 and 22 being less than
1/5 the distance between their bases; the anterior and marginal setae are longer;
seta Z5 is at least 2.\ times as long as seta 75.
Sternal shield granular, presternal area reticulate. SL=^i-82[i; SW= 103-133^;
SL/SW= 0-4-07. Sternal setae subequal in length, first pair slightly shorter than
the others ; metasternal setae about half as long as third pair of sternal setae.
Genital shield flask-shaped, moderately widened behind the genital setae, granular
in appearance, with a faint reticulate pattern. GW^ i 03-162^; GW2= 75-105 pi;
GWj/GW2= 1-2-1-5. Genital setae relatively short, not more than half GW2.
Anal shield triangular, distinctly wider than long. AL=52-94fx; AW=83-ii4(j.;
AL/AW=o-5~o-7. Paranal setae level approximately with middle of anus, or
slightly anterior, about i£ times as long as postanal seta.
Integument of idiosoma bears about 20 pairs of simple ventral and marginal
setae, of which 3 pairs border the genital shield. Metapodal plates elongate, length
to width about 6 : i.
Chaetotaxy of legs normal. Posterior seta on coxa I is i|-i| times as long as
the anterior seta. Anterior seta on coxa III about twice as long and thick as
posterior seta. Length of tarsus IV about 4 times the width at the base.
MALE (fig. 136) : Fixed digit of chelicera with a slender pilus dentilis as in the
female. Dorsal shield as in the female except that the centrally situated setae are
relatively longer. Holoventral shield expanded behind coxae IV, bearing 23 setae,
5 pairs lying in the region between coxae IV and the anus. Integument of iosoma
with about 13 pairs of setae. Chaetotaxy of legs as in female.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE 5. lot. 71
HOSTS AND LOCALITIES: Bathyergus suillus (Schreber) from Strandtfontein, Cape
Town (types in Dr. C. D. Radford's collection and in M.R.A.C.), and from Tulbagh
District, Western Cape Province (S.A.I.M.R.).
Cryptomys hottentotus (Lesson) from Bloemfontein, Orange Free State, and
Pietermaritzburg, Natal (S.A.I.M.R. and B.M.N.H.).
136
FIGS. 136-138. Androlaelaps scapularis (Berlese), venter of male (Fig. 136).
Androlaelaps spatuliformis (Lavoipierre), venter (Fig. 137) and chelicera (Fig. 138) of
male.
Cryptomys holosericeus (Wagner) from Pietermaritzburg, Natal (S.A.I.M.R.).
NOTE: The specimens from Bathyergus suillus are larger than those from the
Cryptomys species, but there are no reliable features which could be used to separate
these groups. The types of /. bathyergus, H. eloffi and H. natalensis have been
compared with drawings of the type of H. scapularis Berlese made by Dr. F.
Gorirossi (Keegan, 1956) and it is believed that they are all conspecific. The deep
posterior concavity of the sternal shield, indicated by Zumpt & Till (1953) in their
drawing of H. eloffi, appears to be the result of over-clearing.
72 W. M. TILL
Androlaelaps spatuliformis (Lavoipierre)
Haemolaelaps spatuliformis Lavoipierre, 1956, Ann. trop. Med. Parasit. 50 : 293, fig. 2.
FEMALE (figs. 139-140) : Deutosternum provided with at least 4 rows of 2-3 small
teeth, 5th and 6th rows very indistinct. Chelae about 40^ long; pilus dentilis and
arthrodial filaments not clearly visible in the one specimen available for examination.
139
140
FIGS. 139-140.
Androlaelaps spatuliformis (Lavoipierre), female,
dorsum (Fig. 140).
Venter (Fig. 139);
Dorsal shield oval; DL=525(x; DW=36o(j.; DL/DW=i-4- Several of the setae
are missing from the specimen examined, but the chaetotaxy appears to follow the
38- or 39-pair pattern. Seta px$ cannot be distinguished in this specimen. Seta
Z5 is more than twice as long as seta /5.
Sternal shield has a few very indistinct reticulations; presternal area weakly
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 73
reticulate. Sternal and metasternal setae subequal in length; first pair of sternal
setae inserted on presternal area. SL= 94-96^; SW= 140-1 43^; SL/SW=O7.
Genital shield spatulate in shape, very slightly widened behind genital setae,
then tapering gradually; GW1=ioo[j.; GW2=97;j.. Genital setae relatively short,
about i GW2.
Anal shield slightly longer than broad; AL=ioo-ii2[x; AW=9i^.. Paranal setae
inserted near posterior margin of anus, about f as long as postanal seta.
Integument of idiosoma bears about 19 pairs of ventral and marginal setae.
Metapodal plates provided with a pronounced medial protrusion.
Chaetotaxy of legs normal except for genu IV which has only 9 instead of 10 setae.
Posterior seta on coxa I about i| times as long as anterior seta.
MALE (figs. 137-138) : Structure of chelicera indicated in fig. 138 ; pilus dentilis not
visible. Dorsal shield as in female; seta px$ present. DL= 404-409^; DW=266^;
DL/DW=i-5. Holoventral shield expanded behind coxae IV, bearing 21 setae,
excluding the first pair of sternal setae which is situated on the presternal area.
Chaetotaxy of legs as in the female. Anterior and posterior setae of coxae I
approximately equal in length and thickness. Femur II has one stout ventral
spine and tarsus II has two stout ventral spines.
HOST AND LOCALITY : Anomaluris derbianus Gray from Kumba, British Cameroons.
One female paratype received on loan from Dr. M. M. J. Lavoipierre, and two males
from the type host and locality received from the Liverpool School of Tropical
Medicine, by courtesy of Miss M. Johnson.
Androlaelaps spreo (Zumpt & Till)
Haemolaelaps spreo Zumpt & Till, 1956, Z. Parasitenk. 17 : 290, fig. 9.
FEMALE (figs. 141-143): Androlaelaps spreo is very closely allied to A. dasymys,
differing from it mainly in the structure of the chelicerae. The pilus dentilis is
different in shape and the arthrodial filaments are subequal in length. The chelae
are 54~56pL long.
The dorsal shield bears 39 pairs of setae as well as 4-6 accessory setae between the
/ series. All the setae are relatively long, the lengths of setae 24 and Z2 being at
least 1-5 times the distance between their bases. The marginal setae are slightly
longer and thicker than those on the median part of the shield and are barbed.
Seta Z5 is 17-1-8 times as long as seta /5. DL=849~936[x; DW=525~6o4fz;
DL/DW=i-6.
Sternal shield reticulate; SL= 114-140^; SW=i66-i75[A; SL/SW=07~o-8.
Genital shield slightly widened posteriorly, with a pronounced reticular pattern
extending to the anterior margin. GW^ 158-175^; GW2= 114-1 23^; GWJ
GW2=i-3-i-4. Genital setae relatively long, slightly more than 2/3 GW2.
Anal shield approximately as long as broad and with a reticular pattern; AL=
140-162^ ; AW= 140-158(4. Anus situated at slightly more than its length from the
anterior margin of the shield; paranal setae about 4/5 as long as postanal seta.
Integument bears about 33 pairs of ventral and marginal setae, the latter being
74
W. M. TILL
longer and thicker than the former and provided with barbs. Metapodal plates
oval, 1-5-1-8 times as long as broad.
Chaetotaxy of legs normal. Posterior seta on coxa I slightly longer and thicker
than anterior seta. Tarsus II has three stout ventral bristles but no blunt spines.
Trochanter IV has a stout dorsal bristle. Length of tarsus IV 6|~7| times width
at base.
141
142
FIGS. 141-143. Androlaelaps spreo (Zumpt & Till), female. Venter (Fig. 141); dorsum
(Fig. 142); chelicera (Fig. 143).
MALE (figs. 144-145) : Chelae as illustrated, pilus dentilis resembling that of female.
Dorsal shield bears 8 accessory setae, otherwise chaetotaxy as in female. DL=
675^; DW=447[A; DL/DW=i-5. Holoventral shield reticulate, expanded, bearing
23 setae. Chaetotaxy of legs as in female.
HOSTS AND LOCALITIES: Spreo bicolor (Gmelin), Springs, Transvaal (type series,
S.A.I.M.R.).
Riparia paludicola (Vieillot), Johannesburg, Transvaal (S.A.I.M.R.).
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 75
Androlaelaps steyni (Till)
Haemolaelaps steyni Till, 1959, /. ent. Soc. S. Afr. 22 : 430, figs.
FEMALE (figs. 148-150) : This species is very closely allied to A. mesopicos. Dorsal
shield relatively shorter and broader than in A. mesopicos and bearing 38 pairs of
setae. DL= 578-648^; DW= 368-420^; DL/DW=i-5-i-6.
Sternal shield relatively shorter than in A. mesopicos. SL= 53-79(1 ; SW=
144
147
146
FIGS. 144-147. Androlaelaps spreo (Zumpt & Till), venter (Fig. 144) and chelicera
(Fig. 145) of male.
Androlaelaps steyni (Till), venter (Fig. 146) and chelicera (Fig. 147) of male.
149-166(1; SL/SW=o-3-o-5. Genital shield similar to that of A. mesopicos. Anal
shield triangular, with a straight anterior margin; AL= 96-105(0.; AW— 105-114^;
AL/ AW = 0-9-1-0. Postanal seta more than twice as long as paranal setae.
Coxa III has a greatly thickened anterior spine; tarsus IV has only one stout
terminal spine. Length of tarsus IV about 5 times width at base.
MALE (figs. 146-147) : Chelicera as figured, fixed digit bearing a relatively short,
moderately inflated pilus dentilis. Dorsal shield similar to that of female, but
76
W. M. TILL
with longer marginal setae. DL= 530-534^; DW=3i5-333[j.; DL/DW= 1-6-1-7.
Coxa III has a stout anterior spine as in the female.
HOSTS AND LOCALITIES: Lybius torquatus (Dumont), Mabelikwa, Transvaal (type
series, S.A.I.M.R. and B.M.N.H.).
Trachyphonus vaillanti Ranzani, near Johannesburg, Transvaal (S.A.I.M.R.).
Lamprotornis nitens (Linnaeus), Rustenburg, Transvaal (S.A.I.M.R.).
FIGS. 148-150. Androlaelaps steyni (Till), female. Venter (Fig. 148) ; dorsum (Fig. 149) ;
ventral view of tip of tarsus IV (Fig. 150).
Androlaelaps suncus sp. nov.
FEMALE (figs. 151-153) : Deutosternum provided with 6 rows of 2-5 teeth. Chelae
about 30[j, long ; movable digit with a pronounced terminal hook which arches over
the tip of the fixed digit ; pilus dentilis inflated, hour-glass-shaped in some positions ;
arthrodial filaments subequal in length.
Dorsal shield oval, reticulate, widest about the level of seta Si. DL= 608-694^;
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 77
DW=385-456{x; DL/DW= 1-5-17. The shield bears the usual 39 pairs of setae
and one accessory seta. Lengths of setae 14 and 22 about 2/3 the distance between
their bases ; seta Z$ nearly twice as long as seta 75. Setae Z$ and 53-55 are barbed,
the others appear to be simple.
Sternal shield reticulate, except in the postero-median third. SL = 90-105^ ;
151 152
FIGS. 151-153. Androlaelaps suncus sp. nov., female. Venter (Fig. 151); dorsum
(Fig. 152); chelicera (Fig. 153).
= 128-148^; SL/SW=o-7. Metasternal setae and first pair of sternal setae
subequal in length, 2nd and 3rd pairs of sternal setae slightly longer.
Genital shield slightly widened behind the genital setae, broadly rounded
posteriorly, with a fairly well-marked, overall, reticular pattern. GW^— 105-120^ ;
GW 2=84-10311; GWj/GW^i-2. Length of genital setae about 3/5 GW2.
Anal shield approximately as long as broad; AL— 103-120^; AW= 106-115^.
Anus slightly more than its length from the anterior margin. Paranal setae level
approximately with middle of anus, about 5/6 as long as postanal seta.
78 W. M. TILL
Integument of idiosoma bears about 23 pairs of ventral and marginal setae, the
latter being barbed. Metapodal plates elongate, length to width about 4:1.
Chaetotaxy of legs normal. Posterior seta on coxa I is i| times as long as anterior
seta. Length of tarsus IV about 5 times its width at the base.
MALE: Not known.
HOSTS AND LOCALITIES: Suncus varius (Smuts) from Tibedi, Basutoland,
9 October, 1953 (4 females).
Crocidura sp. from Pilgrim's Rest, Transvaal, 14 July 1951 (one female not included
in the type series).
Holotype and one female paratype in the collection of the South African
Institute for Medical Research, two female paratypes (1962.6.12.18-19) in the
collection of the British Museum (Natural History).
Androlaelaps tachyoryctes (Radford)
Haemolaelaps tachyoryctes Radford, 1941, Parasitology 33 : 309, figs. 3-4; Keegan, 1956, /.
Egypt, publ. Hlth. Ass. 31 : 236, figs. 19-21.
Hypoaspis (Haemolaelaps} tachyoryctes, Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 71.
FEMALE (figs. 154-157) : Deutosternum provided with 6 rows of 2-4 very small
teeth. Chelae 34-36^ long; pilus dentilis moderately inflated, elongate, tapering to
a point; arthrodial filaments subequal in length.
Dorsal shield granular, with a faint reticulate pattern. DL= 580-627^; DW=
SSS-SS0^ i DL/DW= 1-7-1-8. The shield bears 39 pairs of setae in the type
specimens, but in the other material examined several of the r series are sometimes
off the plate. Most of the setae are relatively short and fine ; setae 24 and Z2 about
half as long as the distance between their bases; seta Z$ at least twice as long as
seta /5 and considerably thicker.
Sternal shield granular, presternal area reticulate; SL=jo-jg\±; SW=i45-i48(x;
SL/SW=o-5-o-6. First pair of sternal setae situated on presternal area, as long as
metasternal setae, 2nd and 3rd pairs of sternal setae longer.
Genital shield granular, slender, tapering behind the genital setae; the length of
the latter about 3/4 GW2. GW^ 76-86^; GW2=8o^.
Anal shield granular, very strongly convex anteriorly. AL=ii4-i22[z; AW=
72-80^; AL/ AW =1-4-1 -6. Anus situated at 1/3 of its length from the anterior
margin. Paranal setae level approximately with middle of anus, about 4/5 as long
as postanal seta and thinner.
Integument of idiosoma bears 23 pairs of ventral and marginal setae, two pairs of
ventral setae bordering the genital shield. Metapodal plates small, nearly triangular
in shape, length to width about 2:1.
Chaetotaxy of legs normal. Setae on coxa I subequal in length. Tarsus II
bears 3 stout ventral setae and 3 stout, blunt, apical setae. Tarsus III has 3, and
tarsus IV has 4 stout, rather blunt, apical setae. Length of tarsus IV slightly less
than 4 times its width at the base.
MALE (figs. 158-159) : Fixed digit of chelicera bears a slightly inflated, tapering
pilus dentilis. Chaetotaxy of dorsal shield as in the female, but all the setae are
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat.
79
156
157
158
FIGS. 154-159. Androlaelaps tachyoryctes (Radford). Venter (Fig. 154), dorsum (Fig.
155) and chelicera (Fig. 156) of female; ventral view of tarsus II of female (Fig. 157).
Venter (Fig. 158) and chelicera (Fig. 159) of male.
8o W. M. TILL
situated on the shield. DL— 475-49450.; DW= 290-304(0.; DL/DW— 1-6-1-7.
Holoventral shield greatly expanded behind 4th pair of coxae and bears 23 setae,
5 pairs in the region between coxae IV and the anus. Integument of idiosoma
bears 13 pairs of setae. Chaetotaxy of legs as in female.
HOSTS AND LOCALITIES: Tachyoryctes splendens (Riippell) from several localities
in Kenya (types in B.M.N.H., other material presented by Miss J. B. Walker to
S.A.I.M.R.).
Lemniscomys striatus (Linnaeus), Njoro, Rift Valley Province, Kenya (Keegan,
1956).
Androlaelaps taterae (Zumpt & Patterson)
Hypoaspis (Haemolaelaps) taterae Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 83, fig. 6.
Haemolaelaps taterae, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 244.
FEMALE (figs. 160-162): A. taterae is very closely allied to A. villosissimus,
differing from it chiefly in being more densely haired. This is particularly noticeable
in the region between the genital and anal shields, where there are about 13 setae,
and in the area between the genital shield and the meiapodal plates.
The chelae of A. taterae are 44-48(0, long; pilus dentilis strongly inflated basally,
terminal half slender and sharply bent; arthrodial filaments variable in size, the
longest being about twice the average length.
Dorsal shield faintly reticulate and has a dense overall covering of setae.
DL= 813-900(0,; DW= 480-550(0.; DL/DW= 1-6-1-7.
Sternal shield reticulate, sharply demarcated from the reticulate and granular
presternal area. SL= 128-148(0,; SW= 166-175(0,; SL/SW=o-8-o-9. Metasternal
setae nearly as long as first pair of sternal setae ; the latter are barbed.
Genital shield with an overall reticular and granular pattern, slightly widened
posteriorly. GWX= 150-162(0.; GW2= 135-140(0.; GW1/GW2=i-i-i-2. Length of
genital setae about 3/5 GW2.
Anal shield reticulate; AL= 152-162(0,; AW= 124-138(0,; AL/AW= 1-1-1-2. Anus
situated at approximately its length from the anterior margin. Paranal setae level
with middle of anus, about 4/5 as long as postanal seta. Metapodal plates roughly
oval, length about if — 2\ times the width.
Genu III bears 10 setae, tibia III bears 9, otherwise the chaetotaxy of the legs
follows the normal pattern for the genus. Posterior seta on coxa I slightly longer
than anterior seta; many leg setae barbed. Length of tarsus IV is 9 times its width
at the base.
MALE (figs. 163-164) : Chelicera as figured, the fixed digit bearing a pilus dentilis
similar to that of the female. Dorsal shield and integument as in female; DL=
599(0.; DW=359(o,; DL/DW=i-7. Holoventral shield reticulate, bearing numerous
setae on the posterior portion.
HOSTS AND LOCALITIES: Widely distributed in Southern Africa on Tatera afra
(Gray). Single specimens have been recorded from Rattus natalensis (Smith),
Rattus chrysophilus (de Winton) and Otomys irroratus (Brants) (S.A.I.M.R.,
B.M.N.H., and M.R.A.C.).
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lal.
162
164
163
FIGS. 160—164. Androlaelaps taterae (Zumpt & Patterson). Venter (Fig. 160), dorsum
(Fig. 161) and chelicera (Fig. 162) of female. Venter (Fig. 163) and chelicera (Fig. 164)
of male.
ZOOL. 10, i
82
W. M. TILL
Androlaelaps tateronis (Radford)
I schnolaelaps -tateronis Radford, 1939, Parasitology 31 : 247, fig. 4.
Hypoaspis (Haemolaelaps} tateronis, Zumpt & Patterson, 1951, /. ent. Soc. S. Afr. 14 : 71.
Haemolaelaps tateronis, Zumpt & Till, 1953, Ann. Inst. Med. trap. Lisboa 10 : 242.
Haemolaelaps radfordi Keegan, 1956, /. Egypt, publ. Hlth. Ass. 31 : 241, figs. 24-25 (syn. nov.}.
FEMALE (figs. 165-167) : Deutosternum provided with 6 rows of 2-4 teeth. Chelae
165
166
FIGS. 165-167.
Androlaelaps tateronis (Radford), female. Venter (Fig. 165); dorsum
(Fig. 166); chelicera (Fig. 167).
about 36^ long ; pilus dentilis inflated basally, terminal portion slender and curved ;
arthrodial filaments subequal in length.
Dorsal shield oval, reticulate. DL= 656-664^; DW= 400-409^; DL/DW=
1-6-1-7. The shield bears 39 pairs of setae of moderate length; lengths of setae 44
and Z2 approximately equal to the distance between their bases; seta Z$, the longest
on the shield, is about twice as long as seta 75.
Sternal shield reticulate, sharply demarcated from granular and reticulate
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. hit. 83
presternal area. SL= 101-116(0.; SW= 136-1 37^; SL/SW^o-y-o-g. Metasternal
seta about as long as first sternal seta, second and third setae slightly longer.
Genital shield narrow, tapering behind genital setae, with a reticulate pattern
which extends to the anterior margin. GW2= 106-113^; genital setae extend
beyond bases of first pair of flanking setae, their length about 3/4 GW2.
Anal shield pear-shaped. AL= 124-139(0,; AW= 96-106(0. ; AL/AW=i-3. Anus
situated at about its length from the anterior margin; paranal setae level approxi-
mately with middle of anus, about as long as postanal seta.
Integument of idiosoma bears 21 pairs of setae, of which 2 pairs flank the genital
shield, a third pair being placed between the genital and anal shields. Some of the
longer setae have very fine barbs. Metapodal plates slender, length to width
about 6 : i.
Tibia III has 9 setae; the number on tibia II is not clear in all the specimens
examined, but is either 9 or 10. Posterior seta on coxa I twice as long as anterior
seta and considerably thicker. Anterior seta on coxa II and both setae on coxa III
thickened. Three of the apical setae on tarsus II stout and spine-like, one of them
blunt. Length of tarsus IV 6 times the width at the base.
MALE: Not known.
HOSTS AND LOCALITIES: Tatera valida (Bocage), Gulu District, Uganda (syntype
of /. tateronis in B.M.N.H.).
" Rat ", Yei, Sudan (paratype of H. radfordi in B.M.N.H.).
Tatera nigricauda Peters, Lali Hills, Kenya (S.A.I.M.R.).
Unidentified rodents from Uganda (B.M.N.H.) and from the Cameroons (I.R.S.C.).
Androlaelaps tauffliebi sp. nov.
FEMALE (figs. 168-170) : Deutosternum provided with 6 rows of 3-4 teeth; anterior
rostral seta twice as long as capitular seta. Chelae 24(0. long ; movable digit divided
into two parts, a ventral (medial) part which is edentate, except for the terminal hook,
and a dorsal (external) part which is broad and dentate. Fixed digit weakly
sclerotized, bearing a flagellar pilus dentilis. About 4 of the arthrodia] filaments
conspicuously long and stout.
Dorsal shield widest about level of seta ?% with a granular appearance and very
faint indications of a reticular pattern. Laterally it has a distinct double outline.
DL=532-56o[o,; DW= 400-418(0.; DL/DW=i-3-i-4. The shield bears the usual
39 pairs of setae, *3-*'5, 22-2:3, /i-/5» and px2-p%3 being reduced to microsetae.
Seta Z5 five times as long as seta /5. In all the specimens examined the anterior
margin of the shield is tucked under, so that setae si become marginal in position
and ii and n are ventral.
Sternal shield about 3 times as wide as long, deeply concave posteriorly, granular
in appearance. SL= 43-48^; SW= 138-143(0.; SL/SW=o-3O-o-35. Sternal seta III
nearly i| times as long as sternal seta I ; metasternal seta short and fine, about half
as long as sternal seta I.
Genital shield large, broadly rounded posteriorly, granular. GWX= 142-157(0.;
GW2=i20-i33(x; GW1/GW2=i-2-i-3. Genital setae short, about 1/3 GW2.
W. M. TILL
170
171
FIGS. 168-172. Androlaelaps taujfiiebi sp. nov. Venter (Fig. 168); dorsum (Fig. 169)
and chelicera (Fig. 170) of female. Venter (Fig. 171) and chelicera (Fig. 172) of male.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lal. 85
Anal shield with some granulation and striations laterally. AL=77~79[x;
AW=9i-98[j.; AL/AW=o-8-o-9. Anus situated at about half its length from the
anterior margin. Paranal setae inserted near anterior third of anus, about the
same length as the postanal seta.
Integument of idiosoma bears about 20 pairs of setae, some of the marginal ones
having fine barbs. Metapodal plates very long and slender, length about 10 times
width.
Chaetotaxy of legs normal, setae simple, without any marked modifications.
Posterior seta on coxa I approximately i| times as long as anterior seta and slightly
thicker. Length of tarsus IV about 6 times its width at the base.
MALE (figs. 171-172) : Deutosternum and gnathosomal setae as in female.
Chelicera provided with an elongate spermadactyl ; minute teeth present at tip of
movable digit; fixed digit reduced, membranous. Total length of chela about
Dorsal shield as in female. DL=4i8[x; DW=295(i; DL/DW=i-4. Sterni-
ventral shield separated from anal shield, granular in appearance, with faint indica-
tions of reticulation, and bearing 8 pairs of setae. Anal shield approximately as
long as broad, otherwise similar to that of female. Metapodal plates relatively
broader than in the female.
Chaetotaxy of legs as in female, except that three setae on leg II are strongly
inflated, namely, the upper, antero- ventral seta on femur II, upper ant ero- ventral
seta on tibia II, and postero-ventral seta on tarsus II.
HOST AND LOCALITY: Seven females and one male from Cryptomys mechowi
Peters, Lac Calundo, Angola, 6 January, 1955 (from the collection of Dr. R. Taufflieb,
I.R.S.C., leg. A. de Barros Machado).
Holotype (1962.2.12.20), allotype (1962.6.12.21) and two female paratypes
(1962.6.12.22-23) in the collection of the British Museum (Natural History) ; two
female paratypes presented to Dr. R. Taufflieb, Institut de Recherches Scientifiques
au Congo ; two female paratypes presented to the Dundo Museum, Angola.
Androlaelaps theseus Zumpt
Hypoaspis (Androlaelaps) theseus Zumpt, 1950, Parasitology 40 : 301, figs. 3-4.
Turkiella theseus, Zumpt & Till, 1953, Ann. Inst. Med. trap. Lisboa 10 : 225.
Androlaelaps theseus, Cooreman, 1954, Ann. Mus. Congo Beige Tervuren, Zool. i : 163, figs. 1-4.
FEMALE (figs. 173-175) : Deutosternum provided with 6 rows each of 4-8 teeth.
Chelae io6-io8[j. long; pilus dentilis slender, arthrodial filaments subequal.
Dorsal shield reticulate and granular. DL= 1350-1570^; DW=iioo-i235fji;
DL/DW=i-2-i-3. The shield bears 40 pairs of setae, an additional pair being
present in the ax position. The setal lengths vary in mites from different hosts.
Seta r-L distinctly longer than seta ii ; setae 14 and zz slightly longer than the distance
between their bases; seta Z$ at least i| times as long as seta 75.
Sternal shield granular and reticular, anterior and posterior margins concave.
SL=i43-i70|ji ; SW=266-2?6y. ; SL/SW=o-5-o-6. First sternal seta reaches almost
to base of third; second and third sternal and metasternal setae at least i| times as
long as first sternal seta,
86
W. M. TILL
Genital shield granular, slender, very slightly widened behind genital setae,
then tapering posteriorly. GWX= 205-247^; GW2= 190-220^; GW1/GW2=i-i-i-2.
Length of genital setae exceeds the width of the shield at the level of their bases.
Anal shield reticulate and granular. AL= 180-205^; AW= 209-238(1; AL/AW—
o-8-i-o. Anus situated at approximately its length from the anterior margin.
Paranal setae level with posterior half of anus, about f as long as postanal seta.
Integument of idiosoma bears about 42 pairs of ventral and marginal setae, many
of which appear to be barbed. Metapodal plates broadly oval, length approximately
ij times breadth.
174
FIGS. 173-175. Androlaelaps theseus Zumpt, female. Venter (Fig. 173); dorsum
(Fig. 174); ventral view of femur, genu and tibia of leg II (Fig. 175).
Chaetotaxy of legs normal. Anterior and posterior setae on coxae I subequal in
length, the posterior one slightly thicker. Spur on second femur relatively small,
its length being about 1/5 the width of the femur. Inner and outer ventral setae
on tibia II subequal in length and thickness ; inner and outer ventral setae on genu II
both simple. Tarsus II has 3 stout, blunt, terminal spines. Length of tarsus IV
about 7 times its width at the base.
MALE (fig. 176-177) : Gnathosoma as in female ; fixed digit of chelicera elongate,
membranous, bearing a slender pilus dentilis.
Dorsal shield resembles that of female; DL= 1283-1302^; DW= 920-940^;
DL/DW=i-4- Holoventral shield strongly reticulate and also granular, not
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lot. 87
expanded behind coxae IV, bearing 23 setae of which 10 (6+4) are situated in the
preanal region behind coxae IV.
Chaetotaxy of legs as in female, but with inner ventral seta on tibia II stouter,
and inner ventral seta on genu II shorter and stouter.
HOSTS AND LOCALITIES: Widely distributed in Southern Africa on Tatera afra
(Gray), and found occasionally on Rattus natalensis (Smith), Rattus chrysophilus
FIGS. 176-178. Androlaelaps theseus Zumpt, venter (Fig. 176) and chelicera (Fig. 177) of male.
Androlaelaps villosissimus (Berlese), venter female (Fig. 178).
(De Winton), Acomys cahirinus (Desmarest), Saccostomus campestris Peters and
Steatomys pratensis Peters (Zumpt & Till, 1961). Recorded from Tatera afra (Gray)
in the Belgian Congo (Cooreman, 1954). Material in S.A.I.M.R. and B.M.N.H.
Androlaelaps villosissimus (Berlese)
Hypoaspis (Haemolaelaps) villosissimus Berlese, 1918, Redia 13 : 128; Zumpt & Patterson, 1951,
/. ent. Soc. S. Afr. 14 : 72, fig. 5.
Haemolaelaps villosissimus, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 244; Keegan,
, Trans, Amer, micr, Soc. 75 : 314, figs. 1-2,
88 W. M. TILL
FEMALE (fig. 178) : Deutosternum provided with 6 rows of 3-4 small teeth.
Chelae 40^ long; pilus dentilis and arthrodial filaments indistinct.
Dorsal shield with only faint indications of a reticular pattern. DL=7i8-744(z;
DW=429~455[x; DL/DW= 1-6-1-7. The shield has a dense overall covering of
setae which completely obscures any pattern. Most of the setae appear to have
fine barbs.
Sternal shield reticulate, sharply demarcated from presternal area. SL=
no-i2O(ji; SW= 153-162^; SL/SW= 0-7-0-8. Metasternal setae as long as first
pair of sternal setae.
Genital shield granular, with an overall reticular pattern. GW^ 140-144^;
GW2=n8-i23(ji; GW1/GW2=i-i-i-2. Length of genital seta about half GW2.
Anal shield reticulate anteriorly and antero-laterally and also granular in appear-
ance. AL= 105-1 i4[x; AW=96-ioOfj,; AL/ AW == 1-1-1-2. Anus less than its
length from the anterior margin of the shield. Paranal setae level with middle of
anus, about 4/5 as long as postanal seta.
Integument of idiosoma bears numerous setae, 8 or 9 being situated between
the genital and anal shields. The ventrally placed setae appear to be simple,
whereas the marginal ones are barbed. Metapodal plates oval, length about twice
width.
Genu III bears 10 setae and tibia III bears 9; otherwise the chaetotaxy follows
the normal Androlaelaps pattern. Posterior seta of coxa I slightly longer than
anterior seta; many leg setae barbed. Length of tarsus IV 10 times width at base.
MALE: Not known.
HOSTS AND LOCALITIES : Saccostomus campestris Peters, Zululand (Berlese, 1918)
and Bulawayo, Southern Rhodesia (S.A.I.M.R.). Unidentified rodent from the
Cameroons (I.R.S.C.).
Androlaelaps walker ae sp. nov.
FEMALE (figs. 179-181) : The hypostomal processes form two pairs of prominent
brushes, one ventral in position and the other dorsal. Deutosternum provided
with 6 rows of 2-3 teeth. Chelae about 48^ long; pilus dentilis slender; arthrodial
filaments subequal in length.
Dorsal shield oval, reticulate, showing pronounced hypertrichy which extends
anterior to setae 23 and which is not confined to the median part of the shield.
The basic chaetotactic pattern is thus partially obscured. DL= 760-780^;
DW=475-494|x; DL/DW=i«5-i-6.
Sternal shield reticulate, anterior margin not sharply defined. SL=95-iiO(j.;
SW= 133-142(0.; SL/SW= 0-7-0-8. Sternal and metasternal setae subequal in
length.
Genital shield flask-shaped, with a reticular pattern extending to the anterior
flap. GWj= 133-142^; GW2=86-io5[j.; GW1/GW2=i-4-i-5. Genital setae as
long as first pair of sternal setae and approximately equal to GW2.
Anal shield arched anteriorly. AL= 142-152^ ; AW= 103-110^; AL/AW=i-4.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 89
Anus situated at approximately its length from the anterior margin of the shield.
Paranal setae level with middle of anus, i| times as long as the barbed postanal
seta.
179
180
FIGS. 179-181. Androlaelaps walkevae sp. nov., female. Venter (Fig. 170); dorsum
(Fig. 1 80); chelicera (Fig. 181).
Integument of idiosoma provided with numerous setae, nearly all of which appear
to be barbed. Metapodal plates oval or irregular in shape, length to width not
more than 3:1.
Chaetotaxy of legs normal except that tibia I has 14 and tibia III has 9 setae.
Tarsus II has 3 stout terminal and 3 stout ventral bristles, and many of the leg
go
W. M. TILL
setae are barbed. Posterior seta of coxa I slighty longer than anterior seta. Length
of tarsus IV 7 times width at base.
MALE: Not known.
HOST AND LOCALITY: Sixty-three females from Tachyoryctes splendens (Riippell),
Muguga North, Kenya, 6 July, 1954, presented by Miss J. B. Walker, East African
Veterinary Research Organization, to the South African Institute for Medical
Research. Holotype and 38 paratypes in the collection of the S.A. Institute for
FIGS. 182-184. Androlaelaps wilkini (Till), female. Venter (Fig. 182); dorsum (Fig.
183); chelicera (Fig. 184).
Medical Research, 25 paratypes (1962.6.12.23-33) in the collection of the British
Museum (Natural History).
Androlaelaps wilkini (Till)
Haemolaelaps wilkini Till, 1959, J. ent. Soc. S. Afr. 22 : 432, figs.
FEMALE (figs. 182-184) : Deutosternum provided with 6 rows of 3-4 small teeth.
Chelae about 40^ long; pilus dentilis small, inflated; arthrodial filaments subequal
in length.
Dorsal shield oval, reticulate; DL= 709-770^; DW=455-5oo(x; DL/DW=i'5-i-6.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 91
The shield bears 37 pairs of setae, ^4 and r6 being inserted on the adjacent integu-
ment. Marginal setae of the r and 5 series barbed, the S setae being at least twice
as long and thick as those of the / series. Seta Z$ sinuous, 2-^-3 times as long as
seta /5. Setae on the median part of the shield (i series posterior to 13, J series,
Z2-Z3, Zi-Z4, p%2-px3) subequal in length, setae 24 and 22 slightly more than half
as long as the distance between their bases.
Sternal shield and presternal area both reticulate. SL—^o-ygyi; SW= 145-158^;
SL/SW=o-5- First pair of sternal setae slightly shorter than 2nd and 3rd pairs and
situated on presternal area. Metasternal setae as long as first pair of sternal setae.
Genital shield relatively slender, very slightly widened behind the genital setae,
lateral margins almost parallel, posterior portion with a reticulate pattern.
GW1=i23-i49[x; GW2=96-ii4n; GW1/GW2=i -1-1-3.
Anal shield pear-shaped, with a reticulate pattern anteriorly and antero-laterally.
AL=i23-i40[x; AW=ii4-i27[o.; AL/AW=i-i-i-2. Paranal setae on a level
between middle and posterior margin of anus, their length not more than 2/3 that
of the postanal seta.
Integument of idiosoma bears 24 pairs of ventral and marginal setae, the longer
ones being provided with fine barbs; posterior terminal pair long and whip-like.
Metapodal plates slender, slightly curved.
Tibia III has 9 setae; anterior and posterior setae on coxa I uniform in length and
thickness. Tarsus II bears two blunt terminal spines; tarsi III and IV also have a
pair of terminal spine-like bristles, but these are more slender and pointed than the
spines on tarsus II. Length of tarsus IV 5 times width at base.
MALE: Not known.
HOSTS AND LOCALITIES: Myrmecocichla formicivora (Vieillot), Debeete, Bechuana-
land (type series, S.A.I.M.R. and B.M.N.H.).
Merops apiaster Linnaeus, Johannesburg, Transvaal (S.A.I.M.R.).
Androlaelaps zulu (Berlese)
Hypoaspis (Haemolaelaps) inops var. zulu Berlese, 1918, Redia 13 : 125.
Haemolaelaps inops var. zulu, Keegan, 1956, /. Egypt, publ. Hlth. Ass. 31 : 234, fig. 18.
FEMALE (figs. 185-187) : Deutosternum provided with 6 rows each of 4 narrow
pointed teeth. Chelae 40(j. long ; pilus dentilis strongly inflated basally, distal third
or fourth slender, with recurved apex; arthrodial filaments very short, subequal in
length.
Dorsal shield reticulate, bearing 39 pairs of setae and 1-3 unpaired accessory setae
between the / series. Setae on median part of shield relatively long, lengths of
setae 14 and 22 about i£ times the distance between their bases. Marginal setae
slightly longer and thicker, with fine barbs. Seta Z5 at least i£ times as long as
seta 75. DL=627-76ofji; DW= 400-494^; DL/DW=i-5-i-6.
Sternal shield reticulate, sharply demarcated from reticulate presternal area.
SL=ioo-io5(x; SW=i28-i52{x; SL/SW=o«7-o-9. Metasternal setae about as long
as first pair of sternal setae.
92 W. M. TILL
Genital shield reticulate, moderately expanded behind genital setae. GWX =
143-170^; GW2= 105-1 14^; GW1/GW2=i-3-i-5. Genital setae relatively short,
slightly more than half GW2.
Anal shield reticulate antero-laterally. AL=g6-i2O[i.; AW= 103-127^; AL/AW
=0-9-1-0. Anus situated less than its length from the anterior margin of the
shield. Paranal setae between middle and posterior end of anus, reaching beyond
base of, and about 4/5 as long as, postanal seta.
185
FIGS. 185-187. Androlaelaps zulu (Berlese), female. Venter
(Fig. 1 86); chelicera (Fig. 187).
(Fig. 185); dorsum
Integument of idiosoma bears about 22 pairs of marginal and ventral setae, of
which 3 pairs border the genital shield. Marginal setae coarser than the truly
ventral ones, and provided with fine barbs. Outer metapodal plates variable,
usually cigar-shaped, length 2^-3-J- times the width,
ETHIOPIAN MITES OF THE GENUS A N D R O L A E L . IP S BE RLES E 5. tat. 93
Chaetotaxy of legs normal. Anterior seta on coxa I about 4/5 as long as posterior
seta. Length of tarsus IV at least 6 times width at base.
MALE: Not known.
HOSTS AND LOCALITIES: Lemniscomys griselda (Thomas), Rattus natalensis (Smith)
and Otomys irroratus (Brants) from the Transvaal (S.A.I.M.R.).
188
189
FIGS. 188-190. Androlaelaps zuluen&is Zumpt, female. Venter (Fig. 188); dorsum
(Fig. 189); ventral view of femur, genu and tibia of leg II (Fig. 190).
Tatera afra (Gray) from Zululand (S.A.I. M.R.).
(Mites from the above hosts were previously identified by Zumpt & Till (1956) as
forms of Haemolaelaps glasgowi.)
Keegan (1956) has recorded this species from several rodents in Egypt.
94 W. M. TILL
Androlaelaps zuluensis Zumpt
Hypoaspis (Androlaelaps} zuluensis Zumpt, 1950, Parasitology 40 : 300, fig. i.
Turkiella zuluensis, Zumpt & Till, 1953, Ann. Inst. Med. trop. Lisboa 10 : 218, figs. 2-5.
Androlaelaps zuluensis, Keegan, 1956, /. Egypt, publ. Hlth. Ass. 31 : 229, fig. 14.
FEMALE (figs. 188-190) : Deutosternum provided with 6 rows each of 3-4 very
small teeth. Chelae 46^ long; pilus dentilis slender; arthrodial filaments short and
subequal in length.
Dorsal shield reticulate, oval. DL= 656-675^; DW— 403-486^; DL/DW=i-5.
The shield bears 39 pairs of setae as well as 2 or 3 unpaired accessory setae between
the J series. Lengths of setae 24 and Z2 approximately equal to the distance
between their bases; seta Z$ about i| times as long as seta /5. The marginal setae
appear to be barbed.
Sternal shield reticulate, posterior margin almost straight or slightly irregular.
SL=io5[j.; SW=n8-i24[A; SL/SW=O'9. Sternal and metasternal setae all
relatively short, a little more than half the length of the sternal shield.
Genital shield with a pattern of striations on its posterior portion; very long,
extending almost to the anal shield. GW1=iyo[i; GW2= 118-124^; GWJGWg
= 1-4. Genital setae very short, less than half GW2.
Anal shield with a reticulate pattern; AL= 105-109^; AW=io5[j.; AL/AW=i-o.
Anus situated at less than its length from the anterior margin of the shield. Paranal
setae near posterior margin of anus; postanal seta broken in both the specimens
examined.
Integument of idiosoma bears 20-22 pairs of marginal and ventral setae. Meta-
podal plates slender, length 3-6 times the breadth.
Chaetotaxy of legs normal. Anterior seta on coxa I slightly shorter than
posterior seta. Spur on femur II rather slender, thumb-like, its length a little less
than half the width of the femur at its base. Inner ventral seta on genu II short,
spur-like; inner ventral seta on tibia II considerably stouter than outer one; tarsus II
terminates in pointed setae only. Length of tarsus IV about 8 times its width at
the base.
MALE (fig. 191) : Deutosternum with 6 rows of 2-5 small teeth. Spermadactyl
6o(A long, fixed digit bears a slender pilus dentilis.
Dorsal shield similar to that of female; DL=52Opt. (approx.) ; DW=36o[u.; DL/
DW=i-4- Holo ventral shield reticulate, expanded behind coxae IV, bearing 23
setae, 5 pairs of setae being situated in the preanal region behind coxae IV.
Chaetotaxy of legs similar to that of female, except that inner ventral setae on genu II
and tibia II are more slender.
HOSTS AND LOCALITIES : Rattus namaquensis (Smith) from Abercorn Pont, Zululand
(holotype in S.A.I.M.R.), and from Kapps Siding, S. W. Africa (S.A.I. M.R.)
Rattus chrysophilus (De Winton), Southern Africa (Zumpt & Till, 1961).
Petromyscus collinus (Thomas & Hinton), Kamanjab-Ohopoho Road, S.W. Africa
(S.A.I.M.R.).
Galago senegalensis Geoffroy (the figured specimens) from Karamoja, Uganda
(S.A.I.M.R.).
ETHIOPIAN MITES OF THE GENUS AND1WLAELAPS BERLESE s. hit. 95
Keegan (1956) records this species from Arvicanthis niloticus (Desmarest) and Mus
musculus Linnaeus in Egypt.
Androlaelaps zutnpti sp. nov.
FEMALE (figs. 193-195): Deutosternum bears 6 rows of 3-5 very fine teeth.
Chelae about 40^. long; pilus dentilis inflated; one arthrodial filament strikingly
longer than the others ; an extension of the arthrodial membrane between the
chelae is elongate, reaching the tips of the chelae.
191
192
FIGS. 191-192. Androlaelaps zuluensis Zumpt, venter of male (Fig. 191).
Androlaelaps zumpti sp. nov., venter of male (Fig. 192).
Dorsal shield reticulate, widest in posterior half at level of setae Si and 52.
DL= 656-732 [A; DW= 442-494(0. ; DL/DW= 1-3-1-5. The shield bears 39 pairs of
setae ; lengths of setae 24 and 22 a little more than half the distance between their
bases ; length of seta /i is f that of seta Si ; seta Z$ is 2-2 1 times as long as seta 75.
Sternal shield reticulate, sharply demarcated from reticulate and granular
presternal area. SL=86-ii4(j.; SW= 124-133^; SL/SW=o-7-o-9. Metasternal
96
W. M. TILL
and first pair of sternal setae subequal in length, 2nd and 3rd pairs of sternal setae
slightly longer.
Genital shield scarcely widened behind genital setae, tapering posteriorly, with a
well-defined reticular pattern extending over the anterior as well as posterior parts.
193
194
FIGS. 193-195. Androlaelaps zumpti sp. nov., female. Venter (Fig. 193); dorsum
(Fig. 194); chelicera (Fig. 195).
GW1=95-ii4[ji; GW2=95-io5[j.. Genital setae about as long as first pair of sternal
setae and about 2/3 GW2.
Anal shield about 133^ long, 90^ wide; AL/AW=i-5. Anus situated at less than
its length from the anterior margin of the shield. Paranal setae near posterior
margin of anus, about 2/3 as long as postanal seta.
Integument of idiosoma bears about 23 pairs of ventral and marginal setae, those
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 97
near the posterior extremity of the body having fine barbs. Metapodal plates
slender, length at least 5 times the width.
Chaetotaxy of legs normal, anterior seta on coxa I about 4/5 as long as posterior
seta. Trochanter IV bears a stout dorsal seta. Length of tarsus IV about 6 times
its width at the base.
MALE (fig. 192) : Deutosternum bears 6 rows of 4-7 very fine teeth. Fixed digit
of chelicera bears a pilus dentilis similar in shape to that of the female.
Dorsal shield 55O[j. long, 352[A wide; chaetotaxy similar to that of female. Holo-
ventral shield strongly reticulate, bearing 23 setae, 5 pairs being placed in the region
between coxae IV and the anus. Chaetotaxy of legs as in the female.
HOSTS AND LOCALITIES: Holotype (female), allotype and one male and three
female paratypes from Rattus (Aethomys) sp., Richmond, Cape Province, 31 March,
1959, two female paratypes from Rattus (Aethomys) sp., Middelburg, Cape Province,
28 March, 1959. Holotype, one male and two female paratypes in the collection
of the South African Institute for Medical Research; allotype (1962.6.12.34) and
two female paratypes (1962.6.12.35-36) in the collection of the British Museum
(Natural History).
NOTE: Several populations of mites have been examined which are very similar
to A . zumpti, except that the setae of the dorsal shield are longer (lengths of setae
24 and Z2 about i^ times the distance between their bases) and 2-4 accessory setae
are present. These mites, all from the collection of the S.A. Institute for Medical
Research, have been taken from the following hosts and localities:—
Rattus chrysophilus (De Winton), Matopos, S. Rhodesia.
Rattus namaquensis (Smith), Bulawayo, S. Rhodesia; Abercorn Pont, Zululand;
several localities in S.W. Africa.
Rattus paedulcus (Sundevall), Kaokoveld, S.W. Africa.
Otomys unisulcatus Cuvier, Luckhoff, Orange Free State.
Rhabdomys pumilio (Sparrman), Cape Province and S.W. Africa.
Petromyscus collinus (Thomas & Hinton), Kamanjab-Ohopoho, S.W. Africa.
Specimens from Parotomys littledalei Thomas, van Rhynsdorp, Cape Province, are
intermediate between these and the short-haired form, the lengths of setae 24 and Z2
being slightly less than the distance between their bases. For the present, these
will all be regarded as forms of A. zumpti.
HOST-PARASITE LIST.*
CLASS AVES
Order FALCONIFORMES
FAMILY AEGYPIIDAE
Gyps coprotheres (Forster)
Androlaelaps pater soni (Zumpt & Till)
* The host names are the same as those used by Zumpt (1961).
ZOOL 10, I
98 W. M. TILL
Order CORAGIIFORMES
FAMILY MEROPIDAE
Merops apiaster Linnaeus
Androlaelaps wilkini (Till)
FAMILY PHOENIGULIDAE
Phoeniculus purpureus (Miller)
Androlaelaps phoeniculi (Zumpt & Till)
FAMILY CAPITONIDAE
Lybius torquatus (Dumont)
Androlaelaps steyni (Till)
Trachyphonus vaillanti Ranzani
Androlaelaps steyni (Till)
FAMILY PICIDAE
Campethera abingoni (Smith)
Androlaelaps haydocki (Till)
Thripias namaquus (Lichtenstein)
Androlaelaps haydocki (Till)
Mesopicos griseocephalus (Boddaert)
Androlaelaps mesopicos (Radford)
Order PASSERIFORMES
FAMILY TURDIDAE
Myrmecocichla formicivora (Vieillot)
Androlaelaps wilkini (Till)
FAMILY HIRUNDINIDAE
Riparia paludicola (Vieillot)
Androlaelaps spreo (Zumpt & Till)
FAMILY STURNIDAE
Lamprotornis nitens (Linnaeus)
Androlaelaps steyni (Till)
Spreo bicolor (Gmelin)
Androlaelaps spreo (Zumpt & Till)
FAMILY PLOCEIDAE
Spermestes cucullatus Swainson
Androlaelaps congoensis sp. nov.
CLASS MAMMALIA
Order INSEGTIVORA
FAMILY MACROSCELIDIDAE
Elephant Shrew
Androlaelaps arvicanthis Radford
FAMILY SORICIDAE
Suncus varius (Smuts)
Androlaelaps suncus sp. nov.
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lal. 99
Crocidura spec.
Androlaelaps suncus sp. nov.
Crocidura manni Peters
(?) Androlaelaps hystrici (Zumpt & Till)
Order PRIMATES
FAMILY LORISIDAE
Galago alleni Waterhouse
Androlaelaps galagus (Lavoipierre)
Galago senegalensis Geoffrey
A ndrolaelaps zuluensis Zumpt
Order CARNIVORA
FAMILY MUSTELIDAE
Ictonyx striatus (Perry)
Androlaelaps marshalli Berlese
FAMILY VIVERRIDAE
Suricata suricatta (Schreber)
Androlaelaps marshalli Berlese
Order LAGOMORPHA
FAMILY LEPORIDAE
Oryctolagus cuniculus (Linnaeus)
Androlaelaps marshalli Berlese
Order RODENTIA
FAMILY BATHYERGIDAE
Bathyergus suillus (Schreber)
Androlaelaps scapularis (Berlese)
Georychus capensis (Pallas)
Androlaelaps capensis (Hirst)
Androlaelaps cryptomius (Radford)
Androlaelaps georychi sp. nov.
Cryptomys holosericeus (Wagner)
Androlaelaps capensis (Hirst)
Androlaelaps scapularis (Berlese)
Cryptomys hottentotus (Lesson)
Androlaelaps capensis (Hirst)
Androlaelaps marshalli Berlese
Androlaelaps scapularis (Berlese)
Cryptomys mechowi Peters
Androlaelaps tauffliebi sp. nov,
ioo W. M. TILL
FAMILY HYSTRICIDAE
Hystrix cristata Linnaeus
Androlaelaps hy strict (Zumpt & Till)
FAMILY SGIURIDAE
Sciurus carolinensis Gmelin
Androlaelaps casalis (Berlese)
Heliosciurus gatnbianus (Ogilby)
Androlaelaps heliosciuri sp. nov.
Xerus inauris (Zimmermann)
Androlaelaps casalis (Berlese)
FAMILY ANOMALURIDAE
Anomalurus derbianus (Gray)
Androlaelaps spatuliformis (Lavoipierre)
FAMILY PEDETIDAE
Pedetes capensis (Forster)
Androlaelaps marshalli Berlese
FAMILY MUSCARDINIDAE
Claviglis spec.
Androlaelaps ghanensis sp. nov.
FAMILY RHIZOMYIIDAE
Tachyoryctes spec.
Androlaelaps callosus (Berlese)
Tachyoryctes splendens (Riippell)
Androlaelaps marshalli Berlese
Androlaelaps tachyoryctes (Radford)
Androlaelaps walker ae sp. nov.
FAMILY MURIDAE
Arvicanthis niloticus (Desmarest)
Androlaelaps arvicanthis Radford
Androlaelaps marshalli Berlese
Androlaelaps murinus (Berlese)
Rattus spec.
Androlaelaps zumpti sp. nov.
Rattus chrysophilus (De Winton)
Androlaelaps dasymys (Radford)
Androlaelaps marshalli Berlese
Androlaelaps later ae (Zumpt & Patterson)
Androlaelaps theseus Zumpt
Androlaelaps zuluensis Zumpt
Androlaelaps zumpti sp. nov.
Rattus morio (Trouessart)
Androlaelaps rhodesiensis (Zumpt & Patterson)
Rattus namaquensis (Smith)
Androlaelaps dasymys (Radford)
Androlaelaps marshalli Berlese
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESK s. hit. 101
Androlaelaps zuluensis Zumpt
Androlaelaps zumpti sp. nov.
Rattus natalensis (Smith)
Androlaelaps arvicanthis Radford
Androlaelaps dasymys (Radford)
Androlaelaps marshalli Berlese
Androlaelaps murinus (Berlese)
Androlaelaps later ae (Zumpt & Patterson)
Androlaelaps iheseus Zumpt
Androlaelaps zulu (Berlese)
Rattus paedulcus (Sundevall)
Androlaelaps marshalli Berlese
Androlaelaps zumpti sp. nov.
Rattus rattus (Linnaeus)
Androlaelaps casalis (Berlese)
Androlaelaps dasymys (Radford)
Androlaelaps rhodesiensis (Zumpt & Patterson)
Rhabdomys pumilio (Sparrman)
Androlaelaps dasymys (Radford)
Androlaelaps glasgowi (Ewing)
Androlaelaps marshalli Berlese
Androlaelaps murinus (Berlese)
Androlaelaps zumpti sp. nov.
Lophuromys flavopunctatus Thomas
Androlaelaps callosus (Berlese)
Androlaelaps murinus (Berlese)
Lophuromys sikapusi (Temminck)
Androlaelaps dasymys (Radford)
Oenomys hypoxanthus (Pucheran)
Androlaelaps murinus (Berlese)
Dasymys incomtus (Sundevall)
Androlaelaps dasymys (Radford)
Androlaelaps murinus (Berlese)
Lemniscomys griselda (Thomas)
Androlaelaps marshalli Berlese
Androlaelaps zulu (Berlese)
Lemniscomys striatus (Linnaeus)
Androlaelaps murinus (Berlese)
Androlaelaps tachyorycles (Radford)
Acomys cahirinus (Desmarest)
Androlaelaps theseus Zumpt
Saccostomus campestris Peters
Androlaelaps marshalli Berlese
Androlaelaps oliffi (Zumpt & Patterson)
Androlaelaps rhodesiensis (Zumpt & Patterson)
W. M. TILL
Androlaelaps theseus Zumpt
Androlaelaps villosissimus (Berlese)
Cricetomys gambianus Waterhouse
Androlaelaps cricetomydis sp. nov.
Androlaelaps galagus (Lavoipierre)
Petromyscus collinus (Thomas & Hinton)
Androlaelaps zuluensis Zumpt
Androlaelaps zumpti sp. nov.
Steatomys pratensis Peters
Androlaelaps marshalli Berlese
Androlaelaps rhodesiensis (Zumpt £ Patterson)
Androlaelaps theseus Zumpt
Otomys spec.
Androlaelaps murinus (Berlese)
Otomys irroratus (Brants)
Androlaelaps dasymys (Radford)
Androlaelaps glasgowi (Ewing)
Androlaelaps murinus (Berlese)
Androlaelaps later ae (Zumpt & Patterson)
Androlaelaps zulu (Berlese)
Otomys saundersiae Roberts
Androlaelaps dasymys (Radford)
Otomys sloggetti (Thomas)
Androlaelaps dasymys (Radford)
Otomys unisulcatus Cuvier
Androlaelaps zumpti sp. nov.
Parotomys brantsi (Smith)
Androlaelaps dasymys (Radford)
Parotomys littledalei Thomas
Androlaelaps dasymys (Radford)
Androlaelaps zumpti sp. nov.
Mystromys albicaudatus (Smith)
Androlaelaps capensis (Hirst)
Androlaelaps dasymys (Radford)
Desmodillus auricularis (Smith
Androlaelaps marshalli Berlese
Androlaelaps oliffi (Zumpt & Patterson)
Gerbillus paeba Smith
Androlaelaps marshalli Berlese
Androlaelaps oliffi (Zumpt & Patterson)
Gerbillus pyramidum Geoffroy
Androlaelaps centrocarpus (Berlese)
Meriones rex Yerbury & Thomas
Androlaelaps longipes (Bregetova)
ETHIOPIAN MITES OF THE GENUS ANDROLAELAPS BERLESE s. lat. 103
Tatera spec.
Androlaelaps arvicanthis Radford
Androlaelaps marshalli Berlese
Tatera afra (Gray)
Androlaelaps dasymys (Radford)
Androlaelaps marshalli Berlese
Androlaelaps oliffi (Zumpt & Patterson)
Androlaelaps taterae (Zumpt & Patterson)
Androlaelaps theseus Zumpt
Androlaelaps zulu (Berlese)
Tatera nigricauda Peters
Androlaelaps marshalli Berlese
Androlaelaps tateronis (Radford)
Tatera valida (Bocage)
Androlaelaps arvicanthis Radford
Androlaelaps marshalli Berlese
Androlaelaps tateronis (Radford)
Taterillus spec.
Androlaelaps centrocarpus (Berlese)
Taterillus emini (Thomas)
Androlaelaps marshalli Berlese
Unidentified rodents
Androlaelaps arvicanthis Radford
Androlaelaps graingeri Zumpt & Patterson
Androlaelaps hirsti (Keegan)
Androlaelaps marshalli Berlese
Androlaelaps tateronis (Radford)
Androlaelaps villosissimus (Berlese)
REFERENCES
BERLESE, A. 1887. In " Acari, Myriopoda et Scorpiones hucusque in Italia reperta." Fasc. XL,
N. 6.
- 1911. Acarorum species novae quindecim. Redia 7 : 429-435.
- 1916. Centuria seconda di Acari nuovi. Redia 12 : 125-177.
- 1918. Centuria quarta di Acari nuovi. Redia 13 : 115-190.
BREGETOVA, N. G. 1952. New species of mites of the genus Haemolaelaps (Gamasoidea
Laelaptidae) parasitizing rodents. Zoo/. Zh. Moscow 31 : 860-882.
- 1956. Gamasid Mites (Gamasoidea). A cad. Sci. U.S.S.R., Moskva 61 : 246pp.
COOREMAN, J. 1954. Acariens du Congo Beige. Ire serie. Ann. Mus. Congo Beige Tervuren
i : 163-168.
COSTA, M. 1961. Mites associated with rodents in Israel. Bull. Brit. Mus. (nat. Hist.) Zool.
8 : 1-70.
DELFINADO, M. U. 1961. Haemolaelaps travisi, a new species of mite from the Philippines
(Laelaptidae : Acarina). Fieldiana, Zoology 44 : 49-51.
FONSECA, F. DA. 1959. Notes d'acarologie. XLI. Haemolaelaps Berlese versus Atricholae-
laps Ewing et Ischnolaelaps Fonseca; Ornithonyssus Samboa versus Bdellonyssns Fonseca.
Mem. Inst. Butantan S. Paulo (1957/58), 28 : 45-54.
104 W. M. TILL
GROKHOVSKAYA, I. M. and NGUEN-HUAN-HOE. 1961. Gamasid mites of North Viet-Nam.
Part 2. Zoo/. Zh. Moscow 40 : 1633-1646.
HIRST, S. 1916. Notes on parasitic Acari. Description of two new African mites of the
family Gamasidae. /. Zool. Res. i : 76-81.
KEEGAN, H. L. 1956. Ectoparasitic laelaptid and dermanyssid mites of Egypt, Kenya and
the Sudan, primarily based on Namru 3 collections, 1948-1953. /. Egypt, publ. Hlth.
Ass. 31 : 199-272.
- i956a. Original illustrations of Haemolaelaps marsupialis Berlese, 1910 and of five
additional Haemolaelaps species described, but not figured by Berlese. Trans. Amer.
micr. Soc. 75 : 314-319.
LAVOIPIERRE, M. M. J. 1955. A description of a new genus of sarcoptiform mites and of three
new species of Acarina parasitic on primates in the British Cameroons. Ann. trop. Med.
Parasit. 49 : 299-307.
- 1956. A description of a new genus and of three new species of mites (Acarina, Para-
sitiformes) parasitic on West African Mammals. Ann. trop. Med. Parasit. 50 : 291-298.
RADFORD, C. D. 1939. Notes on some new species of parasitic mites. Parasitology 31 : 243-
254-
- 1941. Notes on some new species of parasitic mites. Pt. 4. Parasitology 33 : 306-315.
- 1942. New ectoparasitic mites (Acarina) from Uganda. Parasitology 34 : 185-194.
- 1942^. New parasitic mites (Acarina). Parasitology 34 : 295-307.
1944. New parasitic mites from rodents. Parasitology 35 : 161-166.
STRANDTMANN, R. W. 1949. The blood-sucking mites of the genus Haemolaelaps (Acarina :
Laelaptidae) in the United States. /. Parasit. 35 : 325-352.
TAUFFLIEB, R. and MOUCHET, J. 1959. Notes sur les acariens (Acarina; Laelaptidae et
Spinturnicidae) du Cameroun. Ann. Parasit. hum. comp. 34 : 350-353.
TILL, W. M. 1959. Three new Haemolaelaps species (Acarina : Laelaptidae) from birds in
the Ethiopian region, and a redescription of Haemolaelaps mesopicos Radford. /. ent.
Soc. S. Afr. 22 : 423-435.
VITZTHUM, H. 1943. Acarina. In : Bronn, H. G., Klassen und Ordnungen des Tierreiches,
Bd. 5, Abt. IV, Buch 5, Leipzig (Becker & Erler).
ZUMPT, F. 1950. Notes on parasitic mites. I. Some remarks on the family Laelaptidae
(sensu Vitzthum 1943) with descriptions of three new species from African rodents.
Parasitology 40 : 298-303.
— and PATTERSON, P. M. 1950. The Ethiopian species of Hypoaspis subgen. Androlaelaps
Berlese (1903), with description of a new species. S. Afr. J. Med. Sci. 15 : 67-74.
— and PATTERSON, P. M. 1951. Further notes on laelaptid mites parasitic on vertebrates.
A preliminary study to the Ethiopian fauna. /. ent. Soc. S. Afr. 14 : 63-93.
- and TILL, W. 1953. The genera Turkiella nov. ( — Androlaelaps auct.) and Haemolaelaps
in the Ethiopian region, with keys and descriptions of three new species (Acarina :
Laelaptidae). Ann. Inst. Med. trop. Lisboa 10 : 215-249.
— and TILL, W. M. 1956. Notes on Haemolaelaps glasgowi (Ewing) and related forms in
the Ethiopian region, with descriptions of four new species (Acarina : Laelaptidae). Z.
Parasitenk. 17 : 282-291.
- and TILL, W. M. 1958. Notes on the classification and synonymy of gamasid mites
parasitic on vertebrates (Acarina : Mesostigmata) . /. ent. Soc. S. Afr. 21 : 261-273.
and TILL, W. M. 1961. Suborder Mesostigmata. In Zumpt (1961): The arthropod
parasites of vertebrates in Africa south of the Sahara (Ethiopian region). Vol. I
(Chelicerata) . Publ. S.A. Inst. Med. Res. 9 : 17-91.
THE FRANCOLINS,
A STUDY IN SPECIATION
B. P. HALL
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 10 No. 2
LONDON : 1963
THE FRANCOLINS,
A STUDY IN SPECIATION
BY
B. P. HALL
British Museum (Natural History)
Pp. 105-204 ; 2 Text-figures, n Maps
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 10 No. 2
LONDON: 1963
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in jive series, corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 10, No. 2 of the Zoological series.
Trustees of the British Museum 1963
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued March 1963 Price Thirty-one Shillings
THE FRANCOLINS, A STUDY IN SPECIATION
By B. P. HALL
Associate, Department of Z oology > British Museum (Natural History]
CONTENTS
Page
INTRODUCTION ........... 107
THE GENUS . .......... 108
THE GROUPS . . . . . . . . . . .no
SPECIES AND SUBSPECIES . . . . . . . . .no
METHODS . . . . . . . . . . . .112
CHARACTERS . . . . . . . . . . .115
THE SPOTTED GROUP (francolinus, pictus, pintadeanus) .... 123
THE BARE-THROATED GROUP (afer, swainsonii, rufopictus, leucoscepus) . 125
THE MONTANE GROUP (erckelii, ochropectus, castaneicollis , jacksoni, nobilis,
camerunensis, swierstrai) . . . . . . . .131
THE SCALY GROUP (ahantensis, squamatus, griseostriatus) . . .136
THE VERMICULATED GROUP (bicalcaratus, icterorhynchus , clappertoni,
hildebrandti, natalensis, hartlaubi, harwoodi, adspersus, capensis) . . 140
THE STRIATED GROUP (sephaena, streptophorus) . . . . .148
THE RED-WINGED GROUP (psilolaemus, shelleyi, africanus, levaillantoides,
levaillantii, finschi) . . . . . . . . . .152
THE RED-TAILED GROUP (coqui, albogularis, schlegelii] .... 160
SPECIES NOT INCLUDED IN ANY GROUP (lathami, nahani, pondicerianus ,
gularis) ............ 165
CONCLUSIONS . . . . . . . . . . . 168
SUMMARY . . . . . . . . . . . .172
APPENDIX i . Summary of conclusions reached by R. E. Moreau in his paper
" The Vicissitudes of the African Biota in the Late Pleistocene " . . 173
APPENDIX 2. Taxonomic Notes and Synonymy . . . . .175
MAPS ............ 180
REFERENCES ........... 200
INDEX ............ 201
INTRODUCTION
Thirty-nine species of francolin (Francolinus and Pternistis] were recognised in
Peters' Check List of the Birds of the World 2, 1934, of which thirty-four are African
and five Asiatic: one (F. ochropectus Dorst & Jouanin, 1952) has been described
since. I have rearranged Peters' species slightly, now recognising thirty-six in
Africa and five in Asia. As is inevitable a Check List employing conventional
nomenclature gives no indication of the relationship between the species which must
be the basis for any discussion on speciation. The objectives of this paper are
therefore twofold : firstly to set out in the text and on the maps the present relation-
ship between the species. This is not as formidable a task as might be expected
ZOOL. IO, 2 8
io8 B. P. HALL
from the number of species involved, for thirty-seven of the forty-one fall readily
into eight groups, seven of which are African and one Asiatic. (In this paper the
term " group " is used for either a superspecies, in which all members are largely
allopatric, or for a species group which is a rather looser assemblage of related forms
containing some species which are partly sympatric with other members.) These
groups are discussed and mapped separately with particular reference to their
ecology and the relationship of neighbouring or sympatric forms. The four species
that do not fall readily into any group are discussed separately at the end.
Secondly, from the pattern presented by the birds of each group and by their
distribution I have tried to reconstruct the climatic changes, with the resultant
isolating and rejoining of populations, which could have formed this pattern. In
doing so I am fully conscious that the picture we see today gives only fragmentary
clues to the past and that it is impossible to interpret accurately any but the simplest
situations. Nevertheless, even in the most complex groups, I believe there is
some usefulness in making the attempt, for if a series of studies of this nature suggests
changes that from other branches of science are known to have occurred, it may be
possible to give some tentative datings to various steps in speciation.
Throughout the preparation of this paper I have been particularly fortunate in
being able to discuss the problems with R. E. Moreau, who has been preparing
concurrently a paper on " The Vicissitudes of the African Biota in the Late
Pleistocene ". He has allowed me to quote as Appendix i a summary of his conclu-
sions of the major climatic changes that have taken place in Africa in the past
60,000 years. In acknowledging my gratitude to him for this and for invaluable
advice and criticism I must also absolve him from any responsibility for some of
my interpretations of the evolutionary stages, or the conclusions which I have drawn
from them.
I am also deeply indebted to M. P. Stuart Irwin who prepared distribution maps
for me of all the francolins in the National Museum of Southern Rhodesia: also
to Dr. H. Friedmann, C. W. Benson and I. C. J. Galbraith for criticism of the
manuscript. Others to whom I am grateful for loans of specimens, advice, or other
help, include Dr. Dean Amadon, Professor J. Berlioz, P. Blasdale, Mgr. F. O. Cave,
E. M. Cawkell, Dr. James Chapin, P. A. Clancey, Miss M. Court enay-Latimer,
I. H. Dillingham, H. J. de S. Disney, Sir Hugh Elliot, W. V. Harris, M. J. Hollis,
Cdr. A. M. Hughes (who drew figure i), Gen. Sir Gerald Lathbury, C. W. Mackworth-
Praed, Miles Markus, Col. R. Meinertzhagen, Captain C. R. S. Pitman, O. P. M.
Prozesky, Professor H. Schouteden. Rev. Dr. W. Serle, R. H. N. Smithers, B. W. H.
Stronach, M. A. Traylor, Dr. C. Vaurie, Col. Jack Vincent, Professor V. Van Straelen,
John Williams, Dr. J. M. Winterbottom.
THE GENUS
The francolins are recognised as a branch of the sub-family Phasianinae
(Partridges, Quails, Pheasants) which comprises over fifty genera, most of which
are Palaearctic or Asiatic. My interest for this paper is primarily in the African
members of the genus, but the five Asiatic species are included since they provide
THE FRANCOLINS, A STUDY IN SPECIATION 109
some interesting points of comparison. I am not, however, competent to enter
into a discussion on the limitations of the genus in Asia where there are thirty-
eight other genera in the same sub-family some of which such as Rhizothera, Alectoris,
Perdix, Arboricola seem closely related to Francolinus. They have been dis-
tinguished on various structural characters such as the number of tail-feathers, the
size and shape of the bill, or the length of tail, and I have accepted these distinctions
uncritically. It is sufficient to say that the francolins are characterised by a rather
longer, more hooked bill than members of the other genera (except Rhizothera), a
short tail of fourteen feathers, and an upright stance: in the majority of species the
male, at least, is spurred. The range of colours in the plumage is limited to the
shades of ochre, chestnut, brown, black and white produced by melanic pigments,
but the pattern of individual feathers is complex and varied.
The partridge of Madagascar, Margaroperdix, has also some similarity to the
francolins (particularly to the forest francolin F. lathami) in colour and pattern, but
has various structural differences in bill, tail and legs, which make it doubtful if
the two are very closely related. Apart from this in the Ethiopian region there are
no gamebirds that can be regarded as similar to francolins in either structure or
habits, the only others being the quails (Coturnix and Excalfactoria] , the Stone
Partridge (Ptilopachus) — a bird confined to the savanna and steppe belt north of the
Equator — the Congo Peacock (Afropavo), confined to the Congo forest, and the
guinea-fowl family (Numididae).
I would suggest therefore that the closer affinities of Francolinus with Palaearctic
and Asiatic genera indicate that it originated in Asia, becoming separated from the
other game-birds as a species particularly adapted to sub-tropical grasslands, and
that later it spread into southern Europe and Africa. Once in Africa lack of
efficient competition allowed it to spread widely and to exploit varied habitats
which, in Asia, are occupied by other related genera.
If this hypothesis of the origin of Francolinus is accepted it is apparent that
there are factors present in Africa that have encouraged speciation in the com-
paratively recent past (since the emergence of the genus). This is readily acceptable
in view of the climatic changes known to have taken place (see Appendix i).
Fluctuations in humidity and temperature would cause redistribution of the
montane and lowland forests, and the woodlands, savannas and steppes, isolating
and rejoining the populations of their indigenous birds.
Among the francolins various generic divisions have been proposed, the most
commonly accepted being Pternistis for the bare-throated francolins (as in Peters',
1934: 84). However, I agree with White (1952) that the bare throat alone is not a
generic character and in other respects these birds are closer to some of the other
large francolins than are other members of the genus. If a generic division was to
be made I would separate the birds of the first five groups (including the Bare-
Throated Group) which have relatively plain or vermiculated backs, from the
birds of the last three groups with striated or quail-type patterning on the backs,
placing F. gularis with the former and F. pondicerianus and F. lathami with the
latter. However, I regard such a division as unnecessary and leading to difficulties
in respect of the rather atypical species F, nahani.
no B. P. HALL
For a study on speciation francolins have the advantage of being remarkably
sedentary birds for their size, not disposed to fly any great distances, and closely
associated with a particular habitat. Morphologically the bold colours and patterns,
particularly on the underparts, have the advantage of indicating where interbreeding
has or has not taken place between neighbouring forms. Finally their " sporting "
and edible qualities have given them an interest to many people other than
ornithologists so that their ranges and habits are probably rather better known than
those of any comparable group: at the same time these qualities have led to a
persecution of the genus over the last fifty years which, combined with the damage
done by intensive agriculture to the natural habitats, has almost exterminated
species from some areas in which they were common and renders it difficult, if not
impossible, to get further information about the habits of species from some areas
which are now " developed ".
THE GROUPS
Grouping the francolins has presented little difficulty, for the close relationship
of most members is evident from their appearance, ecology and distribution and
has been recognised by previous authors. In a few cases where the relationship is
less obvious this is discussed in the text. I have felt that it facilitates discussion
to give descriptive names to the groups rather than numbers, and the names chosen
indicate an important character common to all members though it has not been
possible in all cases to find a character which is exclusive to the group.
The order in which the groups are listed and discussed is not intended to be syste-
matic except that the first five and the last three appear to form related assemblages.
The groups and the species that comprise them are as follows: —
1. Spotted Group (francolinus , pictus, pintadeanus) .
2. Bare-throated Group (afer, swainsonii, rufopictus, leucoscepus) .
3. Montane Group (erckelii, ochropectus, castaneicollis , jacksoni, nobilis, camerun-
ensis, swierstrai).
4. Scaly Group (ahantensis, squamatus, griseostriatus) .
5. Vermiculated Group (bicalcaratus, icterorhynchus, clappertoni, hildebrandh
natalensis, hartlaubi, harwoodi, adspersus, capensis).
6. Striated Group (sephaena, streptophorus) .
7. Red-winged Group (psilolaemus , shelleyi, africanus, levaillantoides, levaillan-
tii,finschi).
8. Red-tailed Group (coqui, albogularis, schlegelii).
The four species not assigned to any group are the two African forest francolins
lathami and nahani, and the two Asiatic species pondicerianus and gularis.
SPECIES AND SUBSPECIES
In discussing speciation it must always be borne in mind that this is a continuous
divergent progress starting when two populations become isolated from each other.
In isolation each will develop divergent characters, either morphological or ecological,
or both, but there will be a long period during which the two populations are
THE FRANCOLINS, A STUDY IN SPECIATION in
sufficiently alike for free interbreeding to take place if the isolating barrier is removed
(Phase i). If they remain isolated and divergence continues this stage will
gradually merge into one in which the divergent characters become such that
interbreeding is unlikely (but not impossible) if the two populations rejoin (Phase 2).
Again these characters may be ethological, involving different recognition patterns
or voice ; or ecological, so that the two are unlikely to occur on the same ground ;
or could possibly be a difference in breeding seasons. It is in this stage that the
occasional hybrid may be produced in the wild, and in which hybridisation is
possible and frequent under artificial conditions. Finally the two birds diverge
so completely that they are incapable of interbreeding (Phase 3).
In general most populations in Phase i and in the transition period between
Phase i and 2 would be regarded as subspecies and those in Phase 2 and 3 as species,
but there is no line between them and the decision must rest on the assessment of
the taxonomist. This must be based on the degree of divergence and the degree
to which they are known to interbreed. In the francolins members of different
groups may be assumed to have reached, or almost reached, Phase 3 in their
relationship, but the allopatric members of each group may be only in Phases i and 2.
Each group provides a case in which I have had to make an arbitary decision
whether or not to regard neighbouring forms as conspecific. For example, in the
Spotted Group F. francolinus and F. pictus have diverged appreciably, especially
in males, and hybrids are scarce, so I treat them as species. In the Bare-throated
Group the afer block and the cranchii block have also diverged but interbreed
freely, so are treated as conspecific. In the Montane Group F. ochropectus has
diverged morphologically and ecologically from F. erckelii and is isolated, so is
treated as a species, while atrifrons stands in a similar morphological relationship to
castaneicollis but has the same field habits so is considered conspecific. In the
Scaly Group the Niger provides a barrier between F. ahantensis and F. squamatus
so it is not certain if they are capable of interbreeding but they seem to have diverged
sufficiently to justify specific status. In the Vermiculated Group F. hildebrandti
and F. natalensis have diverged, especially in the females, but interbreed to a
limited extent at one point of contact but not apparently at another so are treated
as species. In the Striated Group " rovuma " type birds (with a streaked abdomen)
apparently do not interbreed with " sephaena " types (unstreaked) in the southern
half of the range but hybridise in the north; the divergence is slight, however, and
they are treated as conspecific. In the Red-winged Group the degree of divergence
and the relationship between shelleyi and its three neighbours, africanus, whytei,
and uluensis varies only slightly, but is just sufficient for me to treat africanus as
a species and the other three as conspecific. In the Red-tailed Group the northern
forms of coqui have diverged slightly, both morphologically and ecologically, from
the southern but seem capable of interbreeding so must be considered conspecific.
The foregoing demonstrates the impossibility of defining a species in such con-
ditions. For this reason I have tried to avoid differentiating strongly between
species and subspecies in the discussion, though greater clarity might have been
achieved by setting out the discussion under specific headings (as has been done in
the Red-winged Group where rearrangement of the species made it necessary).
H2 B. P. HALL
On the other hand I have endeavoured to make a clear distinction between those
subspecies which I term " potential species " — namely those which are either
(a) isolated at present, or (6) must be assumed to have been isolated at some period,
since they show a degree of divergence from their neighbours which is too great
to be attributable to local ecological factors — and those subspecies whose divergence
can be attributed to ecological factors and which, as a rule, form part of a cline.
The characters and ranges of " potential species " are denned equally with those of
full species, but other subspecies are noted summarily in smaller print in a section
in each group headed " Other variation ", together with any significant local or
individual variation. On the maps as much information as possible has been shown
on the ranges and relationships of species and subspecies, but this has necessarily
been dependent on the complexity of each map. However, I do not consider it
practical or desirable to recognise numerous steps in a cline, or slight local variations,
and have relegated to the synonymy many names given to such intermediates and
micro-populations. There is more justification in naming isolated populations,
however small, such as some of the montane ones, and in these cases I have been
guided by the degree of divergence shown. For instance, I regard names as desirable
for such highly divergent small populations as the montane francolins found on the
Plateau du Day in (French) Somaliland (ochropectus) and at Mega in southern
Abyssinia (atrifrons), but not for others in the same group in other mountains of
southern Abyssinia which differ only slightly from each other. To avoid en-
cumbering the main discussion with irrelevant detail, notes on taxonomy and
synonymy are confined to an appendix.
In the parts of the paper dealing with evolution the ancestral stock of present
species or of more than one species is referred to with the prefix " proto " (e.g.
proto-coqui and pToto-albogularisjschlegelii) : if clarity demands, the name is followed
by " subsp." or " sp.". The name of the author and date of publication are insert-
ed only for those subspecies not listed by Peters (1934).
METHODS
The extensive collection of francolins in the British Museum has formed the basis
for this study, supplemented by loans of critical specimens from other museums.
All the African specimens examined and all the authentic records I have been able
to trace in literature have been plotted over the Vegetation Map of Africa (Oxford
1959), as many species as possible being entered on a single sheet. This has enabled
me to make a direct comparison of the range of any species with the vegetational
belts and with the range of other species, whether in the same group or not. The
Asiatic species have been plotted with accuracy only in the few critical areas, and
for this the distribution maps of Indian birds prepared by Whistler, but never
published, have been of great assistance. The maps of the groups have been compiled
from these key maps. In a distribution map it is always difficult to know to what
extent the range should be shown as continuous between actual collecting points,
and in this respect the maps differ. For the Asiatic species I have shown the
THE FRANCOLINS, A STUDY IN SPECIATION 113
limits of the range only: for the African species I have, as a general rule, shaded
only areas from which specimens have been obtained, though this may give a rather
distorted picture from parts of the country which are least known ornithologically :
in a few cases, most notably for the Bare-throated Francolins (Map 2), I have felt
that greater clarity in presentation could be achieved by shading the whole area
within the limits of the range: this has been done only with species which, from
their known habitat preferences, could be expected to have a reasonably continuous
distribution within a vegetation belt.
The morphological discussion is based on specimens examined personally unless
otherwise stated. For the field notes I have used the standard regional works as a
basis but these have been immeasurably enriched by the numerous conversations
and letters that I have exchanged with the many field workers whose names appear
in the acknowledgments. Where these workers have been responsible for specialised
information I have tried to give the credit in the text, but they are responsible also
for much of the general information which is incorporated without acknowledgment.
I have found it difficult when discussing ecology and habitat to be consistent
in the descriptive terms used, especially in correlating the parts of southern Africa
with which I am familiar with those of the north for which I have to rely on the
descriptions of others. As far as possible I have followed the terminology employed
by the Vegetation Map of Africa (Map n). In this the vegetational types which
mainly concern the francolins are the Woodlands, Savannas and Steppes (Types
16-25). My own experience in southern Africa and the works of such specialists
in ecology as Benson and Stuart Irwin show the importance of the distinction
between the woodlands in which acacia is rare or absent (the " brachystegia wood-
lands " or " myombo " — Types 18 and 19) and the acacia country, whether it is
termed acacia " woodland ", " savanna " or " steppe ". This distinction applies
not only to the birds of the actual woodlands but also to those of the grasslands
and savannas (i.e. grasslands with scattered trees) with which the woods are inter-
spersed, many birds being strictly confined to either the brachystegia belt or to
acacia. (In the southern and eastern francolins there are, in fact, several species
indigenous to acacia, such as F. sephaena, F. leucoscepus, F. rufopictus, F. adspersus
and F. levaillantoides , but none which is wholly indigenous to the woodland belt,
though the potential species F. s. shelleyi and F. s. whytei and the Bare-throated
F. afer nearly qualify.) I have therefore made a clear distinction in the discussions
between the brachystegia belt and acacia country and throughout the paper the
term " woodland " is used exclusively for types 17-19 unless " acacia woodland " is
specifically stated.
North of the equatorial forest the woodlands are apparently similar in character
to the brachystegia woodlands though composed largely of different species of trees
and less continuous (Type 17). I am told also that, because of this discontinuity,
the transition from the woodland belt to the acacia is less clearly defined. It is
to be expected that there are therefore less clear-cut ecological distinctions in the
birds, but this is not easy to determine from published works, the term " savanna "
being widely used by different authors without clear definition. I have therefore
been less precise in discussing the habitat of northern francolins. (From the maps it
H4 B- P- HALL
appears that F. clappertoni and F. coqui spinetorum may be indigenous to acacia, and
F. schlegelii, F. albogularis and -F. icterorhynchus to the woodland belt.)
The distinction between acacia "woodlands", "savannas" and "steppes" is
largely one of aridity and the transition between the types is necessarily gradual.
It will be appreciated from the foregoing that savannas are found in both the
woodland and acacia belts, but they are necessarily different in character. Where
the term is used without qualification it should be plain from the context which belt
is under discussion.
In attempting to interpret in terms of evolution the pattern presented in each
group by divergence, ecology and ranges, I have started in each case with the
distributional maps. From these I have tried to find in each group the factor that
limits the ranges. More often than not the answer lies in the vegetation map, and
the range of a group, species or potential species will be found to coincide with the
limits of a vegetation belt, this coincidence being particularly common where
acacia savanna or steppe changes to woodland. The presence or absence of moun-
tains, hills or rivers is obviously significant in other cases, but here it must be
remembered that, while the reason for the present discontinuity in the range of a
species may be self-evident, the range must have been continuous in the compara-
tively recent past. In one case, the Red-winged Group, the limits of ranges of
species coincide not so much with the vegetational belts as with the isohyets of total
annual rainfall. In another, the Striated Group, the isolated areas from which one
species (F. streptophorus) is found seem to have nothing in common except that they
are the two ends of a ridge of higher country. In several cases where there seems
no ecological reason for the limits of a species' range the bird is replaced by a member
of another group. From facts such as these I have tried to assess in what con-
ditions each group would be most likely to thrive at the expense of other groups,
and, conversely, what conditions would be sufficiently unfavourable to divide it
into isolated populations from which the species developed. In postulating in
each group successive climatic changes to account for the isolating and rejoining
of the various popualtions I have been guided solely by the pattern presented by
the birds themselves, and not by any consideration of what climatic changes are
known from other evidence to have occurred, except that I have not, I hope, postu-
lated any changes which are, on this evidence, outside the bounds of possibility.
However, in the final section of the paper I have tentatively tried to correlate some
of the climatic eras postulated with those known, as listed in Appendix i.
In the discussion on evolution two assumptions are frequently made which are
not necessarily correct but which seem sufficiently probable to justify their use as
a basis for hypothesis. The first is the premise that the degree of divergence shown
by two isolated forms can be correlated with the length of isolation. This, of course,
is not necessarily true, but it can, I think, be accepted that in similar circumstance
there is a likelihood that speciation will proceed at approximately the same pace;
also that a comparatively long interval must elapse for considerable divergence to
take place in isolates. The corollary premise that isolates that have not diverged
extensively have not been long separated is more open to doubt, and examples
(such as the forest owl Phodilus prigoginei] can be quoted of populations that must
THE FRANCOLINS, A STUDY IN SPECIATION 115
have been long isolated but show little divergence. But I regard these as
exceptional.
The second assumption is that the present species and potential species originated
in some part or parts of their present range. This is a premise that is the more
questionable in the case of species which are not members of superspecies. However,
in the francolins these are exceptional, most of the species being part of a super-
species in which all members are allopatric and which has a more or less continuous
distribution over a large part of Africa. In these cases it is highly improbable that
the origin of any member was in the territory of the present neighbouring forms,
although all may have been outside the present range of the group (i.e. when vegeta-
tion belts were pushed north or south).
In such a contingency each proto-species may be presumed to have held approxi-
mately the same position in regard to its neighbours as it does at present.
CHARACTERS
The study of the francolins in species groups has served to demonstrate that many
characters which might be considered as guides to relationship show variable
degrees of uniformity in each group. For example, in the Bare-throated Group
the extent of bare skin on the face and throat is absolutely constant, and can be
regarded as a " group character ", whereas in other groups closely related species
may have a variable amount of bare skin. It seems useful therefore to discuss some
of these characters separately at the start in order to obtain an understanding of
their importance. A summary of some of these characters is set out in Table i.
SIZE
Little emphasis has been put on size or proportions in this paper, for though there
is considerable individual variation there is reasonable uniformity in most related
species. In the Red-winged Group the long-billed and short-billed populations of
F. shelleyi provide the only example of marked proportional variation. On the
whole general variation in size follows Bergmann's Law in relation to altitude and
latitude, equatorial and lowland birds usually being smaller, but there are several
inconsistencies. It is perhaps worth noting especially the incidence (for which I
cannot account) of exceptionally small birds on the coasts of Kenya and southern
Angola, and, to a lesser extent, Natal, since this feature is found in other genera
besides francolins.
PLUMAGE
(a) Feather patterns. The complexity of the feather patterns has already been
referred to. A number of examples are illustrated below which serve to define some
of the descriptive terms I have used, and which have been arranged in sequences to
indicate some of the transitional stages between one pattern and another. This is
a subject which deserves fuller treatment than I can give it here and much of the
discussion by Harrison in his paper " The incidence and origin of spotted patterns
in the Estrildidae " (awaiting publication in the Ibis) is relevant to the francolins.
It is sufficient here to emphasise that comparatively small changes in feather pattern,
B. P. HALL
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particularly at the tip of the feathers, may greatly alter the appearance of a bird.
(Compare the great differences in the general appearance of the breasts of a female
schlegelii and a male coqui, and in the underparts of a male and female pintadeanus
with the small differences in feathers 2 and i, and 5 and 3 of Figure i.)
(b) Underparts. The greatest variety of colour and pattern throughout the genus
is found on the underparts (and to a lesser degree on the mantle) and considerable
variation is found between closely related birds. The most striking example is
illustrated by Bowen (1930) and shows the difference between the subspecies of
8
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16
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Fig. i. Feather patterns.
i. Barred (c? coqui, breast). 2. Barred with inverted triangle at tip ($ schlegelii,
breast). 3. Transition, barred to spotted ($ pintadeanus, breast). 4. Transition,
barred to spotted (<$ shelleyi, breast). 5. Transition, barred to spotted (<$ pintadeanus,
breast). 6. Spotted (
138 B. P. HALL
is a richer red-brown in the centres of the feathers both above and below, and has
very few white streaks on the abdomen; there is some black in the lores and eye-
stripe. The colour, but not the extent of the red-brown patterning in the mantle,
is matched in a few individuals of schuetti from various parts of its range, but none
approaches this colour below.
The third species of the Scaly Group, F. griseostriatus, is confined to the strip of
rich vegetation with vestigial patches of forest which is associated with the escarp-
ment in western Angola. It is the most distinct member of the group having the
leathers of the mantle and wing-coverts chestnut broadly edged with grey, and the
rest of the upper parts faintly vermiculated, as in F. s. squamatus and F. ahantensis,
but paler: below the feathers of the breast and flanks are chestnut edged greyish
or creamy buff and the abdomen is plain creamy buff. While the coloration is
brighter and more contrasting, the pattern of the feathers of griseostriatus is
fundamentally the same as in the other two species. The bill is mainly blackish
on the upper mandible with a bright red base, and below is orange-red : the legs are
orange-red, and the three males examined have only a single spur.
EVOLUTION
The chief features of the distributional pattern of the Scaly Group are the two
specific divisions which separate ahantensis and griseostriatus from squamatus;
the vast areas in which squamatus exhibits no significant variation : and finally the
indications of speciation in the eastern populations. The fact that the ranges of
F. ahantensis, and F. s. squamatus coincide so closely with the limits of the main
blocks of lowland forest suggest strongly that glades and clearings in this type of
forest must be regarded as the natural habitat for the group, and that the populations
found outside the perimeter of the forest are relicts from a period in which the forest
extended to the extreme limits of the range of the Group. The occurrence of these
populations in montane forest, up to at least 9,500 ft. on some mountains, and the
rapid adaptation of the birds in Kenya to cultivations indicate, however, that their
requirements are not rigid as long as sufficient cover is available. A wide spread of
montane forest might, therefore, not be wholly disastrous for the Scaly Group, though
they could not expect to compete entirely successfully with members of the Montane
Group in these conditions. (It is perhaps significant that the highest altitudes at
which Scaly Francolins have been found are on Mount Elgon, which has apparently
no representative of the Montane Group.)
It seems likely, therefore, that prolonged dry eras have been the chief factors in
speciation. In some such periods (possibly coincidental) early in the history of the
group proto-ahantensis could have been isolated from proto-squamatus by a belt of
savanna stretching to the coast between the Niger and the Volta, and proto-
griseostriastus isolated along the escarpment of Angola, where moister conditions
prevailed, while the interior of the country was dry (Hall, 19606) . With a return
of wetter conditions in West Africa proto-ahantensis spread eastwards. The lack
of divergence between populations on either side of the present gap in the forest east
of the Volta lends support to the conclusions cited by Moreau that the gap between
THE FRANCOLINS, A STUDY IN SPECIATION 139
the two blocks of forest was formerly further east in the neighbourhood of the Niger
delta. The Niger itself at present lies between the two species, and since there can
be little temptation for sedentary birds such as francolins to fly across a broad
river it can be regarded as a barrier sufficiently formidable to inhibit interbreeding.
It is possible that the Lower Congo could have acted as a similar barrier between
proto-squamatus and pitoto-griseostriatus if in a wet era the escarpment forests of
Angola were reunited with the forest of Gabon and Cabinda.
In F. squamatus the remarkable lack of variation in the whole species argues that
the last spread of forest which linked the present outlying populations was of
comparatively recent date. If degree of divergence can be accepted as a guide to
time then it can be postulated that the earliest populations to be isolated by the
retreat of the lowland forest would be those of Nyasaland and Tanganyika: but in
this instance especially this precept must be posed very tentatively for the isolated
populations concerned have also become adapted to rather different conditions at
higher altitudes which might accelerate divergence. Since the most geographically
remote of the remaining populations, those of the Jebel Marra (known from one
specimen only) and southern Abyssinia, do not differ significantly from typical
schuetti in north-eastern Angola and the southern Congo, the forest may have
persisted longest in north-eastern Africa.
Finally the distinction and the fairly sharp transition between squamatus and
schuetti on the perimeter of the Congo forest merits attention. It suggests that at
some time western and eastern birds were divided (presumably in a dry era) and on
reuniting the eastern birds tended to favour rather more open habitat on the
forest fringes, and the western birds clearings in the deep forest.
A summary of the evolution postulated is as follows :—
Stage i. The group developing in lowland forest, and probably extending to East
Africa.
Stage 2. In a dry era proto-ahantensis in the Upper Guinea forest becomes cut off
from proto-squamatus in the Lower Guinea forest somewhere in the region of the
Niger. P-roto-griseostriatus in the escarpment forests of Angola also becomes cut
off from proto-squamatus.
Stage 3. In a wetter, more favourable, era, the group spreads again but no inter-
breeding takes place between the incipient species, the barriers formed by the
Niger and Lower Congo possibly reinforcing any tendency to genetic isolation that
has developed while they have been separated.
Stage 4. A dry era divides the Lower Guinea forest into a western block, in which
proto-squamatus subsp. develops and an eastern block in which the birds become
adapted to slightly less humid conditions. These eastern birds, proto-schuetti,
keeping more to the forest fringes when the two are reunited.
Stage 5. A wide spread of lowland forest extends the range of proto-ahantensis to
the Gambia, and proto-squamatus sp. to embrace all the outlying localities at which
it is found at present (Map 4A).
Stage 6. With the retreat of the forest, populations become isolated first in Nyasa-
land, proto-doni, and Tanganyika, proto-uzungwensis and proto-usambarae, then on
the mountains of the Kenya/Tanganyika border, and finally in the Sudan, Uganda
I40
B. P. HALL
and Abyssinia. These isolated populations survive in isolated patches of lowland
and riverine forest, and also become adapted to montane forest up to high altitudes,
especially where there is no competition from members of the Montane Group, and,
in some areas, to cultivations.
In West Africa populations of ahantensis become isolated in the Gambia and
Portuguese Guinea and the main block of the species is divided by a break in the
forest in the region of the Volta.
THE VERMICULATED GROUP (MAP 5)
(F. Ucalcaratus, F. icterorhynchus, F. clappertoni, F. hildebrandti, F. natalensis, F.
hartlaubi, F. harwoodi, F. adspersus, F. capensis)
RANGE AND CHARACTERS
The Vermiculated Group is the most widespread of all the African groups. I
regard it as consisting of a superspecies (containing six allopatric species) and
three related species. The superspecies has a more or less continuous distribution
from Senegal to Eritrea and southwards to Natal with outlying populations in
Morocco and in the mountains of South West Africa and southern Angola. Of
the three extraneous species one (F. harwoodi} is confined to a small area of Abyssinia
almost encircled by the superspecies: the second (F. adspersus) replaces the super-
species in northern Bechuanaland but is partly sympatric with the outlying
population in South West Africa: the third (F. capensis] is completely isolated in
the extreme south.
The ecology of the group is variable and will be discussed in detail under the
individual species but it can be said in general that from West Africa to Abyssinia
and Uganda the members of the superspecies (F. bicalcaratus, icterorhynchus and
clappertoni) are birds of grasslands and cultivations in the woodland, acacia savanna
and steppe belts. However, in east and southern Africa this habitat is largely
occupied by members of other groups, in particular F. afer of the Bare-throated
Group (Chapin 1932: 714), and it appears that members of the Vermiculated Group
have been driven into more rocky country on hillsides and into thickets in river
valleys and the distribution is consequently more broken (compare Maps 2 and 5).
In choosing the term " vermiculated " as the descriptive name for this group I
have given emphasis to the most striking character which the majority of the
component forms have in common although in some subspecies of F. clappertoni
the vermiculations are largely or wholly replaced by extensive U-patterning. All
members have brown or grey-brown heads, backs, wings and tail, with lighter
vermiculations and/or V- and U-patterning, and sometimes some rufous mottling:
the lores are black or blackish and most forms have a white eye-stripe : the under-
parts are patterned with dark brown (and sometimes chestnut or maroon) on a
whitish or cream background, except in the females of the two sexually dimorphic
forms which are plain orange-ochre below: the colours of the bill and legs, and the
colour and extent of bare facial skin, is variable and so is the number of spurs (see
Table i).
THE FRANCOLINS, A STUDY IN SPECIATION 141
SPECIES AND POTENTIAL SPECIES
The superspecies consists of six clearly differentiated forms which are usually
regarded as species, though it can be argued that two of these, F. hildebrandti and
F. natalensis, could be treated as conspecific. The three northern members,
F. bicalcaratus, F. icterorhynchus and F. clappertoni, form a homogeneous assemblage
occupying grasslands and savannas of various types. F. bicalcaratus is the repre-
sentative in West Africa, being common in the acacia and woodland belts from
Senegal to the Cameroons, extending also into cleared parts of the Upper Guinea
forest. There is also an isolated population in Morocco. Above, it is faintly
vermiculated with extensive V- or U-patterning on the mantle and wing-coverts:
below, each feather is cream with some chestnut on the edges and with a black drop-
shaped centre that is broken by small cream " windows " (Fig. i, 15), giving a
partly streaked and partly spotted effect: the bill and legs are greenish, and two
fairly well-developed spurs are common in the males.
The Morocco population (ayesha) is not conspicuously different, though isolated
by 1,500 miles, but may be slightly more rufous and with rather smaller " windows "
in the patterning below. Other variation in the species is clinal and will be
discussed later.
In the Cameroons, at about longitude I5°E., F. bicalcaratus is replaced abruptly
(but not apparently along any ecological boundary) by a plainer bird, F.
icterorhynchus, which is more vermiculated and less V-patterned above, and which is
irregularly V-patterned, blotched or spotted with dark brown below: the bill and
legs differ from bicalcaratus in being mainly orange, and there is a small patch of
yellow skin round the eye. Its range extends eastwards through the woodland belt
(Map n, types 16 and 17) to Uganda, where a few specimens are found which
have some chestnut streaks on the flanks (as in the type of "F. ugandensis) ." In
the acacia belt to the north (Map n, type 20) F. clappertoni replaces F. bicalcaratus
between northern Nigeria and Lake Chad, and replaces F. icterorhynchus in the Sudan.
It is common also in the Nile and Blue Nile valleys and on grassy hillsides of western
Abyssinia and Eritrea below 7,000 ft. (above which it is replaced by the montane
francolin, F. erckelii). Western birds (F. c. clappertoni) are very similar to F.
icterorhynchus above but a more orange brown, while below they resemble F.
bicalcaratus except that there are no " windows " in the dark feather centres and the
feathers are edged with maroon rather than chestnut. In the Nile valley and southern
Sudan birds become darker and greyer, less vermiculated, but heavily U-patterned
above and more heavily patterned below on a whiter ground. In the Blue Nile
valley and eastwards birds lack vermiculations entirely but are clearly and regularly
V-patterned above and on the breast, giving a lanceolate appearance, and they
lack the moustachial streak which is present in the western birds.
A single specimen collected at " Ngeem ", Lake Chad (possibly Nguigmi on the
north-west edge of Lake Chad), the type of "F. tschadensis ", is very like some of the
specimens of F. clappertoni from the Nile valley but the underparts are more buffy
and lack any maroon markings, and it is rather less heavily patterned on the back.
Neumann suggested (c/Bannerman, i : 327) that it is a hybrid between F. clappertoni
and F. icterorhynchus. Since F. icterorhynchus does not now apparently occur as
i42 B. P. HALT.
far north as this, and since this specimen is rather more heavily patterned than the
northern populations of icterorhynchus and clappertoni it might perhaps be better
regarded as an aberrant specimen that indicates the close relationship of the two
species.
East and south of Lake Victoria F. icterorhynchus is replaced by F. hildebrandti,
which is a bird largely associated with rocky ground and thickets in river valleys
and on hillsides, from sea level to about 8,000 ft., and which is found equally in the
acacia savanna and the brachystegia woodland belts. It is common from central
Kenya south to the Tabora and Dodoma districts of Tanganyika but in the southern
part of Tanganyika and northern Mozambique its distribution is sporadic : however,
it is again common in Nyasaland at varying altitudes, and in the upper Luangwa
valley. The males are very like F. icterorhynchus above, but have more solid,
blotchy, brown markings below: the bill is reddish with a brown culmen and
yellowish base, and the legs reddish. The female is quite different having the
underparts plain orange ochre. The sexual differences are most complete in the
most southerly birds from southern Nyasaland, Mozambique and south-west
Tanganyika (F. h. johnstoni] in which the female has no patterned plumage on either
the upper mantle or the breast, whereas those from most of Kenya (F. h. altumi)
have some patterned " male " plumage on both. Females from the centre of the
range (F. h. hildebrandti) are intermediate, having the patterned feathers mostly
confined to the upper mantle.
F. hildebrandti reaches the southern limits of its range in the Luangwa and Shire
valleys, and is replaced further south by F. natalensis, a bird with apparently the
same ecological preferences. Birds from the Transvaal and Natal and the southern
parts of Southern Rhodesia (F. n. natalensis} differ from hildebrandti in having
the sexes alike, the backs darker brown and more heavily patterned and vermiculated
and having regular double U-patterning below, unlike the brown blotches of the
male hildebrandti. The bill and legs of natalensis are similar to those of hildebrandti
except that there is no black on the culmen and the male has commonly one spur and
the female none, whereas in hildebrandti the male has commonly two (once three)
spurs and the female one or two vestigial spurs. However, the intensity and
regularity of the patterning, particularly in the females, decreases northwards,
showing some approach to hildebrandti. The populations of the northern districts
of Southern Rhodesia, the Zambezi valley and the Luangwa valley (neavei) are
plainer and more rufous above and the females tend to have the underparts washed
with buff, the abdomen plainer and the U-patterning less defined giving a more
spotted or streaked effect, but there is much individual variation.
As far as is known the ranges of F. hildebrandti and F. natalensis only approach
each other in two areas, in the Luangwa valley between latitudes 12° and I3°S and
in the neighbourhood of the Shire/Zambezi confluence. In the Luangwa valley
there is some indication that the two may sometimes interbreed, or have interbred
in the recent past, for some of the specimens of natalensis are more blotched, less
U-patterned below than those of F. n. neavei, and some of the females are washed
with buff below and have plainer abdomens (see Benson and White, 1957 ' 138) :
one male from Jumbe (13° i6'S., 32° 07'E.) lent by the National Museum of
THE FRANCOLINS, STUDY IN SPECIATION 143
Southern Rhodesia is close to F. n. neavei below but has, like hildebrandti, a dusky
ridge to the culmen and two spurs. Nevertheless, there are in this area specimens
typical of the two forms taken from within 50 miles of each other so it is probable
that there is only sporadic interbreeding.
In the Zambezi/Shire area the situation seems rather different. F, natalensis
has been collected from points along the Zambezi as far down as Tambara (60 miles
below Tete) but not as low as the junction with the Shire, and not from the left
bank below the junction of the Mushonganende River, 280 miles above Tete.
Within 40-60 miles of Tambara F. hildebrandti has been collected in southern Nyasa-
land and neighbouring Mozambique at Zobue, " on rocky hillsides " (Vincent 1934:
336), on Mlunganyama hill " at 500 ft. but not at 200 ft." (Benson, C. W. & F. M.,
1948: 3), on Mwananbidzi Hill at 2,500 ft., and near Port Herald in the Shire valley
(Long, 1960: 100), but not as far down the Shire as the confluence. There is no
evidence in this area of interbreeding between F. natalensis neavei and F. hildebrandti
and it has been suggested that here the Zambezi itself, which is as much as two miles
wide when in flood, may be a barrier (Benson, et alia: in press).
The last member of the superspecies, F. hartlaubi, is confined to the mountains of
northern South West Africa and southern Angola and is strictly a bird of the rocks.
It is the smallest member of the group and has distinctive rufous mottling above.
The females are rufous below like those of F. hildebrandti but the male is finely streaked
from chin to abdomen in brown and white. The bill is brownish above and yellow
below and at the base: the legs are yellow with spurs poorly developed but often
with two present vestigially in both sexes.
The first of the three members of the group which are not included in the super-
species is a francolin about which little is known, F. harwoodi. Only males have
been collected and from only three localities, all in the gorges of the Upper Blue
Nile or its tributaries. Of members of the superspecies it resembles F. natalensis
most closely, being similar above but with more defined U-patterning on the nape,
and below having similar double U-patterning, but rather darker and more clearly
defined and extending to the chin, and being absent from the centre of the abdomen,
which is cream. It is distinctive in lacking the white eye-stripe but having a circle
of bare scarlet skin round the eye. The bill is recorded as red but from the appear-
ance of skins probably has a black culmen; the legs are red with two fairly well-
developed spurs. F. harwoodi appears to be cut off from other members of the
group by the presence of the montane francolin F. erckelii on the plateau above the
gorges, in the type of open country which, at lower altitudes, is occupied by F.
clappertoni both to the north and the south.
In southern Africa along the borders of Rhodesia and Bechuanaland the super-
species (represented by F. natalensis) is replaced in similar habitat by F. adspersus,
which ranges westwards through both acacia savanna and steppe (Map u, types 20
and 25) to South West Africa where it overlaps the range of the isolated member
of the superspecies, F. hartlaubi. Here the two forms are found in the same localities
but differ in their ecology and habits, adspersus living in coveys along watercourses
and hartlaubi usually in pairs on the rocks. F. adspersus is a rather different-looking
bird from those members of the group that have been discussed: its appearance is
144 B. P. HALL
grey, the pattern being composed of minute vermiculations on the upper parts
and with broader black-and-white vermiculations over all the underparts. It has
no eyestripe but an area of bare yellow skin round the eye: the bill and legs are
orange-red, the male having usually a single long spur.
The last member of the Vermiculated Group, F. capensis, is isolated in the extreme
south-west. Like many other members of the group it is found in thick cover in
rocky river valleys chiefly in the coastal districts of the Cape. It is not known
from Little Namaqualand (possibly because of the lack of suitable rivers) but has
been recorded from the gorge of the lower Orange River at Assenkjer. It is the
largest member of the group and most like F. natalensis and F. harwoodi in the
predominance of brown and white double U- or double V-patterning. However,
in capensis the patterning is distinctive in being composed of very fine irregular
white lines on most of the back (the rest being vermiculated), and similar on the
underparts but with distinct white shaft streaks, giving a streaky effect: it has no
eye-stripe and no bare skin round the eye : the bill, as in hildebrandti, is red with a
dark culmen, and the legs red, the male having one or two spurs and the female
one.
OTHER VARIATION
In F. bicalcaratus variation is largely clinal, the palest birds being found in the drier savanna
areas and the darkest and most heavily patterned in the cleared parts of the forest areas in
southern Cameroons. Among the dry country birds those from Senegal, Gambia and the
hinterland of Ghana (bicalcaratus) are paler with more rufous heads than those of northern
Nigeria and northern Cameroons (adamauae) . The birds from within the borders of the Upper
Guinea forest from Sierra Leone to Ashanti (thornei) are slightly whiter, less creamy below than
those within the borders of the Lower Guinea forest from south-eastern Nigeria and southern
Cameroons (ogilviegranti) which become increasingly dark eastwards. Birds from Accra, between
the two forest blocks, are closer to bicalcaratus than to thornei: those of most of southern
Nigeria are intermediate between adamauae and ogilviegranti. The isolated ayesha in Morocco
(already discussed) is closest to the paler birds above but has heavier patterning below.
In F. icterorhynchus variation is clinal and ecological, pale, lightly patterned birds being
found in the southern Sudan and in the region of the Albert Nile in northern Uganda (icterorhyn-
chus) : dark, heavily patterned birds are found in the more humid areas from the Oubangi and
Uelle to southern Uganda (dybowskii).
In F. clappertoni the clinal variation is greater than in the two previous species, and features
some changes in pattern that warranted discussion in the main section. West of the Nile
and in the Nile Valley the paler, less patterned clappertoni merges through the intermediate
heuglini to the dark U-patterned gedgii of the southern Sudan. Among the more V-patterned,
lanceolated populations of the east, those of Eritrea, and northern Abyssinia (sharpii) are pale
above (though not as sandy as clappertoni), those of south-western Abyssinia (nigrosquamatus)
are darker and more patterned, and those of the Blue Nile valley (konigseggi) are somewhat
intermediate, closest to sharpii but with a greater suffusion of black markings below.
In the southern populations of F. natalensis there is some variation according to the habitat,
darker, more patterned birds being found in the richer vegetation of Natal, Zululand, and the
Zoutspansberg, and paler birds in the drier savanna country of the Transvaal (see Clancey,
J953 : 59)- (Since the range of the darker birds is discontinuous it is impracticable to distinguish
the two variations by name.)
The available series of F. hartlaubi is inadequate for detailed study, but it would seem that
the populations of southern Angola (hartlaubi) are rather smaller than those of South West
THE FRANCOLINS, A STUDY IN SPECIATION 145
Africa, and that the western birds from the Kaokoveld and Erongo (crypticus Stresemann,
1939) are rather paler than those from the Waterburg and Otavi further east (bradfieldi) .
EVOLUTION
It has already been suggested that the rather different ecology of the northern
members of the Vermiculated Group compared with those from Kenya southwards
may be due to competition with the Bare-throated Group, and this possibility has
considerable bearing on the interpretation of the pattern presented by the ranges
of the various species. I believe, for example, that while the present can be regarded
as generally favourable for the Vermiculated Group as a whole, it is even more
favourable for the Bare-throated Group and that the distribution of the southern
Vermiculated species is heavily restricted because of this, especially in the areas
where alternative habitats to the savanna grasslands are limited.
This would account for the absence of any member of the group from the grass-
lands south of the Congo forest, also for the sporadic distribution in eastern Africa,
and possibly also for the limited ranges of F. natalensis and F. capensis in the
coastal districts of the south-east, where the gap between them is occupied by the
bare-throated F. afer.
In the north-east it is apparent from the isolated population in Morocco that the
group is not at present as widespread as it has been at some time in the not distant
past (to judge by the lack of divergence in the Morocco population). Elsewhere
in the north the range appears to be expanding, for there is good reason to suppose
that F. clappertoni has only recently extended its range in the acacia savanna south
of the Sahara, where it has supplanted F. coqui (see discussion under Red-tailed
Group and Maps 5 & 8), and also has recently extended its range eastwards towards
the Red Sea coast, cutting off the Erkowit population of the montane francolin
(F. e.pentoni] from the main body of the species (compare maps 3 and 5).
From the foregoing it can be assumed that grasslands and scrub in savanna and
woodland are the natural habitat for the group, but it is adaptable : the presence of
several members in areas where the rainfall in under 10 inches a year shows especially
that it can be tolerant of dry conditions if there is adequate cover in watercourses.
It is likely, therefore, that humid eras in which there has been a wide spread of
forest have been less favourable to the group than the dry eras, and that it is chiefly
in these wet eras that the group has been split and speciation has developed in the
isolated populations. The morphological differences between F. adspersus and
other members of the group, combined with the fact that it has diverged sufficiently
to be able to live alongside another member, F. hartlaubi, in complete ecological
segregation, suggest that probably the group developed in the east and south-east
and that pToto-adspersus was isolated earliest from the original stock. This could
conceivably have taken place at the time when a spread of montane forest linked
the Natal forests with those of Southern Rhodesia and Nyasaland, cutting off
proto-adspersus in the Kalahari/South West African region. Pioto-capensis and
proto-harwoodi can be presumed also to have been isolated at an earlier period than
any of the six members of the superspecies, but several factors could have caused
their segregation. Proto-capensis, as suggested, could have been cut off by the
]46 B. P. HALL
intrusion of the bare-throated F. afer. F. harwoodi on the other hand appears to
be a " relict " population that has survived in the Blue Nile Gorges at a period when
the group was overrun on the plateau above. The fact that the high parts of the
plateau immediately surrounding the range of harwoodi are at present occupied by
F. erckelii of the montane group suggests that it was first split from other members
of the group in an era when montane conditions prevailed all over the plateau, as
they do now.
Proto-hartlaubi in South West Africa may also have been cut off at an early date
from the remainder of the superspecies, in which case it must be regarded as a
coincidence that it has diverged along the same very distinctive lines (with marked
sexual dimorphism) as the Tanganyika population, hildebrandti. It seems more
likely that, following the periods in which adspersus, capensis and harwoodi were
isolated, there was a period which favoured the superspecies, during which it became
widespread through the grasslands of the tropics and south to Natal and the
Transvaal. Meanwhile, proto-adspersus had become adapted to rather drier con-
ditions in acacia steppe to the south-west. A subsequent drier era in southern
Africa which spread acacia steppe through the Limpopo and Zambezi valleys could
isolate proto-natalensis from the rest of the superspecies at a time when proto-
hartlaubi and proto-hildebrandti were still linked. A further spread of steppe
northwards bringing competition with proto-adspersus or a spread of the Bare-
throated Group might then eliminate the superspecies from the areas between the
Rhodesias and South West Africa, where there is little alternative habitat. The
mountains of the west, however, would provide sufficient ecological variety for
proto-hartlaubi to find a different habitat and co-exist in South West Africa with
proto-adspersus, and in southern Angola with the bare-throated afer.
The tendency to approach hildebrandti shown by the Zambezi population of
natalensis (neavei) suggests that these birds are either closest to the ancestral stock
of natalensis I hildebrandti or else are the result of a period in which the two had
become reunited after their distinctive characteristics had developed, but before
there was any bar to free interbreeding. However, the lack at present of an inter-
mediate population showing all stages of intergradation between hildebrandti and
natalensis suggests that the ranges of the two were again separated in the not too dis-
tant past : the presence in the Luangwa valley of a few apparent hybrids among other-
wise typical birds indicates that there has been only sporadic interbreeding now that
the ranges again join. Minor fluctuations in climate would serve to bring about this
splitting and reuniting.
The differences between the various populations of hildebrandti are not great
enough to suggest any lengthy periods of isolation, but, combined with the present
discontinuous distribution, indicate a limited amount of splitting and reuniting
since the species developed. The greater amount of "male" plumage exhibited
by the females of hildebrandti in western Kenya (altumi) indicates some approach to
F. icterorhynchus of Uganda, suggesting that the break between them was at a
comparatively recent date.
The three northern members of the superspecies, icterorhynchus, clappertoni and
bicalcaratus form a homogeneous and completely allopatric assemblage from which
THE FRANCOLINS, A STUDY IN SPECIATION 147
it can be postulated that the present extensive range has not been occupied for a
long period, since otherwise climatic fluctuations would have caused complex
splitting and rejoining of populations over such a wide area. To interpret the main
lines of the pattern it is only necessary to visualise one period in which the ancestral
stock of the three members spread westwards from Abyssinia to Senegal, followed
by one unfavourable period in which it was divided into a western (proto-bicalcara-
tus), central (pToto-icterorhynchus) , and eastern (pioto-clappertoni] population.
This period seems likely to have been a humid one in which Lake Chad was extensive
and forest extended from its southern tip southwards, forming a north-south barrier
between the western and central parts of the savanna belt. At the same time it
can be postulated that there would have been sufficient riverine forest along the
Nile to form a second barrier further east, and that forest in Kenya and Uganda cut
off proto-icterorhynchus from proto-hildebrandti in the south. Speciation developed
in the three pockets and was almost complete when dry conditions returned. Proto-
clappertoni became adapted to rather drier conditions than the other two species
and spread eastwards over the Nile north of proto-icterorhynchus. The differences
in pattern between the eastern and western populations of clappertoni suggest
that at some subsequent period the Nile and its tributaries again formed a barrier
between east and west, allowing birds with the distinctive V-patterning of F. c,
sharpii to develop in the north-east. At present it appears that clappertoni is
extending its range westwards, north of the ranges of icterorhynchus and Ucalcaratus,
at the expense of the red-tailed coqui.
In the west it is apparent that there must have been a period when sufficiently
humid conditions prevailed along the coast to allow the francolins to spread to
Morocco. The lack of marked divergence in the Morocco population (ayesha)
suggests that this period was of a more recent date than the humid period postulated
for the isolation of the ancestral stocks of the three northern species.
A summary of the evolution postulated is as follows: —
Stage i. Ancestral stock of the group develops in the east and south-east.
Stage 2. Proto-adspersus isolated in the south, possibly by a spread of montane
forest.
Stage 3. The rest of the group extends northwards to Abyssinia, and widely through
grasslands south of the Congo. Pioto-capensis isolated, perhaps by competition
with the Bare-throated Group.
Stage 4. The group spreads westwards to Senegal but withdraws from the high
plateau of Abyssinia, which is occupied by the Montane Group, leaving a relict
population in the Blue Nile gorges, proto-harwoodi.
Stage 5. (South). Pioto-natalensis isolated, possibly by extension of acacia steppe
in Limpopo or Zambezi valleys bringing competition with proto-adspersus.
Stage 5. (North). Proto-bicalcaratus in the west, proto-icterorhynchus in the centre
and pioto-clappertoni in the east, isolated from each other, probably by a belt of
forest extending northwards to Lake Chad (at a time when its area was more exten-
sive), and by riverine forest in the Nile valley. Pioto-clappertoni becoming adapted
148 B. P. HALL
to drier conditions than the other two species. Forest in Uganda and Kenya
separates proto-hildebrandti from proto-icterorhynchus.
Stage 6. (South). Proto-hartlaubi isolated in the mountains of South West Africa
and Angola, possibly by an invasion of the central areas by proto-adspersus or by
the Bare-throated Group.
Stage 6. (North). Minor fluctuations, allowing pToto-clappertoni first to spread
westwards across the Nile and then splitting into a western and eastern population,
possibly during a return of more humid conditions when the Nile valley again
became a barrier.
Stage 7. (South). Minor fluctuations causing rejoining and re-splitting of proto-
natalensis and proto-hildebrandti, and isolating populations of hildebrandti for short
periods.
Stage 7. (North). A humid period in which proto-bicalcaratus spreads to Morocco,
followed by a drier period in which the Morocco population is isolated.
Stage 8. (South). The ranges of natalensis and hildebrandti rejoin, at least in the
Luangwa valley, and there is sporadic interbreeding.
Stage 8. (North). In the Sudan clappertoni spreads westwards through the drier
acacia belt.
THE STRIATED GROUP (MAP 6)
(F. sephaena, F. streptophoms)
RANGE AND CHARACTERS
The Striated Group is unlike all others in consisting of only two species which are
partly sympatric. The range of the group as a whole is mainly in eastern Africa,
from Somalia to Natal, but extending across to South West Africa and southern
Angola, and with an isolated population in the Cameroons.
Both species are mainly brown above with some chestnut, black and white pattern
in the collar, and white shaft streaks on the mantle: the eye-stripes are white,
edged with black and there is some chestnut on the sides of the face: the throat is
white bordered with chestnut: the underparts are creamy buff with contrasting
maroon, chestnut, or black-and-white patterning on the breast (different in the two
species) and lighter patterning on the flanks and abdomen.
The habitat of the two species is different and will be discussed later.
SPECIES AND POTENTIAL SPECIES
The two species are quite distinct both ecologically and morphologically. F.
streptophoms has a puzzling distribution, being found commonly in the grasslands
and " orchard bush " of northern Uganda and sporadically in north-western Kenya
usually on stony scrub-covered hillsides (Jackson, 1938: 241). In north-western
Tanganyika Dillingham has found it also in clearings of brachystegia. These are
the only areas from which it is known in the east but 1,500 miles to the west a single
specimen was obtained by Bates among grass and rocks between 3,500 and 4,000 ft.,
THE FRANCOLINS, A STUDY IN SPECIATION 149
in the Cameroon highlands 40 miles west of Fumban (on the borders of former
British and French Cameroons).
It is a darker bird than F. sephaena the back being grey-brown with dark brown
patches and with fewer and narrower white shaft-streaks. It has a solid chestnut
collar on the hind neck above a broken black-and-white collar of barred feathers :
there is a considerable amount of chestnut in the facial pattern but only a narrow
band bordering the base of the throat: the feathers of the breast are barred black
and white occasionally with a chestnut tip: the remainder of the underparts is
pale cream marked with some dusky vermiculations and broad blackish streaks on
the flanks : the bill is black with yellow at the base and the legs yellowish with only
a vestigial spur in the males. The sexes are alike. The Cameroon bird is like those
of East Africa but smaller (wing i<$ 145 against 6^ 151-160).
In northern Uganda, where both species are found, Pitman tells me that sephaena
is strictly confined to the driest areas of scrubby thorn country while streptophorus
is in savanna grasslands. Both Jackson (1938: 241) and Dillingham (in litt.} stress
that the call of streptophorus is quite unlike that of any other francolin known to
them, and is a melodious trill or whistle. Dillingham on the other hand found
sephaena' s call closer to other francolins though Vincent, on a label of a bird from
Swaziland, describes it as "a loud piping trill " and Benson tells me he has
found it very variable, Rhodesian birds being hardly recognisable as the same
species as those in Abyssinia. Comparison between the eggs would also be useful
for Pitman confirms Jackson's statement that the eggs of sephaena are exceptionally
hard, and it would be interesting to know if those of streptophorus, at present un-
known, have the same character.
F. sephaena is widespread in the acacia savanna and steppe from eastern Abyssinia
and (British) Somaliland westwards to the Albert Nile, southwards through Kenya
and along the Tankanyika coast to Mozambique and Natal, westwards to South
West Africa and southern Angola. It is never far from water and in the driest
parts of the range is confined to watercourses.
Above it is a rich red-brown with broad white shaft-streaks : the chestnut collar
is broken with white and the patterned feathers of the upper mantle are rich dark
brown or blackish with elliptical white centres : the females are slightly vermiculated.
There is no black on the underparts, the feathers of the breast being cream with
triangular chestnut or maroon markings on the tips : the bill is wholly black and the
legs carmine, the males having a single long sharp spur.
The birds here designated as F. sephaena have sometimes been considered to
belong to two species, for two distinct types are found, one, mainly coastal, which
has fine, drop-shaped maroon streaks on the abdomen, and the other in which the
abdomen may have some dusky patterning but which has no maroon streaks. In
the Shire valley of southern Nyasaland birds of the two types appear to behave as
species being found close to each other without an ecological division, but without
any intergrading : specimens from the southern tip of Lake Nyasa and eastwards
are streaked, and so are those from the lower Shire valley at Chiromo and Tangadzi,
but a specimen from Lengwe, 35 miles upstream, is unstreaked, as are those from
all localities westwards in the Zambezi valley (Benson, C. W. and F. M., 1948: 3).
1.50 B. P. HALL
In northern Tanganyika a similar clear division is found between the two types,
birds from inland, at altitudes above 1,500 ft., being unstreaked, those from the
coastal plain being streaked. However, from the Kenya boundary northwards
there is no clear division either geographically or morphologically: in western Kenya,
southern Abyssinia (Benson, 1945: 391) and throughout Somalia both types are
found, as well as some intermediates with sparse, fine streaks. In (British) Somali-
land streaked birds predominate at lower altitudes near the coast but both types
are found in the highlands and streaked specimens have been found as far inland
as Harar in north-eastern Abyssinia. There is no evidence that the two types
behave here as species and it is more probable that they represent a hybrid population.
(For this population it is convenient to use the name spilogaster, which was given
to a streaked bird from Harar — see Rand, 1950 : 384 while rovuma should be confined
to the unmixed streaked populations from Tanganyika southwards.)
It will be seen from the map that inland birds of the unstreaked type are found
in two blocks from northern Tanganyika northwards and from southern Nyasaland
southwards, the species being absent from inland areas through most of Tanganyika,
Nyasaland and Northern Rhodesia. Birds of the northern block (grantii) have the
patterning on the collar and on the breast more restricted (especially in the females)
than the southern birds. In birds of the southern block the patterning on the
collar is brighter and more extensive and the patterning below extends to the lower
breast: birds from the Transvaal, Natal and extreme southern Mozambique
(Coguno) have the rest of the underparts fairly heavily vermiculated (sephaend),
but northwards through Southern Rhodesia the vermiculations become lighter and
birds from South West Africa and southern Angola across the continent to southern
Nyasaland are appreciably less patterned (zambesiae) .
OTHER VARIATION
Throughout the range of F. sephaena there is considerable local variation in size with
equatorial birds on the whole being smaller. Exceptionally small birds are found on the Kenya
coast and exceptionally large ones in the Upper Zambezi and Chobe valleys. Among birds
without the chestnut streaks on the abdomen there is considerable variation in the amount
of vestigial patterning on the abdomen, some having ill-defined whitish streaks and irregular
dusky U-patterning. On the whole eastern birds in, and bordering, the hybrid zone tend to
be more patterned than western.
In South West Africa some rather greyer birds are found, probably associated with the
limestone pans, but the variation is not constant.
EVOLUTION
The peculiar distribution of F. streptophorus is the major problem set by the
Striated Group. The fact that the eastern and western populations of streptophorus
are not strongly differentiated and yet the country in between appears totally
unsuitable for the species indicates that these are relicts from an era when the group
had a wide distribution north of the Congo forest. Being relicts they may not now
be in their natural habitat and it can, therefore, be reasonably postulated from the
distribution of sephaena, that the natural habitat of the Group is in acacia steppe,
and that proto-streptophorus once occupied this belt from the Sudan westwards,
THE FRANCOLINS, A STUDY IN SPECIATION 151
north of the forest and woodland belts. One thinks usually of these belts as moving
north and south with successive climatic fluctuations, but it is possible to visualise
that the ridge of higher ground that stretches at about latitude 6°N from the
Cameroons eastwards to Uganda may sometimes dislocate regular north-south
movement. It seems possible that, following a dry era in which the belt of acacia
steppe lay as far south as this ridge, the return of more humid conditions and richer
vegetation might come first to the lower country of the Shari basin and Bahr-el-
Ghazal to the north of the ridge, leaving temporarily a pocket of dry country on
the ridge while the main belts moved north. If proto-streptophorus became isolated
in this pocket it could be expected that, as conditions became more humid and less
and less suitable, it would be driven to the only parts where a tolerably dry habitat
could still be found, and one which would be unattractive to the indigenous francolins
of the encroaching woodlands. Such refuges were provided in the higher ground at
the western and eastern extremes of the previous range.
It is possible that streptophoms may yet be found in the country between the
Cameroons and Uganda, but if so could be expected only in some patches of sparse
vegetation. Since so little divergence is shown by the Cameroon bird it can be
postulated that it has been separated from the Uganda population at a comparatively
recent date.
The pattern presented by sephaena points to at least one unfavourable era in
which the species was divided for long enough to allow the streaked and unstreaked
types to develop in isolation, almost to the specific level. The present distribution
shows how small an extension of woodland towards the coast in Tanganyika would
divide the species. In a considerably wetter era with a wide spread of woodland
or forest it is likely that the northern populations would be confined to the coastal
plain of Somalia while the southern populations might develop on the fringes of the
Kalahari. If this was followed by a dry era with a spread of acacia the unstreaked
southern populations might spread northwards up the Luangwa valley, through
Tanganyika and Kenya along the valleys of the rivers which run westwards to the
lakes, and then up the Abyssinian Rift to the Somali highlands.
Meanwhile the streaked northern populations spread southwards, chiefly in the
valleys of the rivers flowing to the sea. It is difficult to account for the fact that the
two types seem to behave as species in the south but mix and interbreed to a limited
extent in the north. It may be that the two came together at a later date in the
south when speciation was further advanced, and it is also possible that in northern
Tanganyika the sharper change of altitude on the edge of the coastal plain may
provide here a barrier that is lacking in Kenya and eastern Somalia (though not in
the mountains of (British) Somaliland).
The present gap in distribution between the unstreaked birds of Kenya and
northern Tanganyika (grantii) and those of the Rhodesias and southern Nyasaland
(zambesiae) is sufficient to account for the divergence between them.
The scattered distribution of sephaena along watercourses in the drier areas would
account for much of the local variation in both size and patterning that is exhibited,
for such populations would be isolated and might be expected to diverge in periods
when there is little suitable vegetation between one river valley and another.
152 B. P. HALL
A summary of the evolution postulated is as follows: —
Stage i. Ancestral stock occupying the acacia belt north, east and south of the
central forest and woodland block.
Stage 2. The stock split into two, one, proto-streptophorus , in the acacia belt north
of the central forest, the other, proto-sephaena, in acacia in the eastern districts from
Somalia to southern Africa. This split probably taking place in a humid era with
a spread of woodland or forest between Uganda and Abyssinia.
Stage 3. F. sephaena split into a northern group, probably confined to lowland in
Somalia, and a southern group, probably on the borders of the Kalahari. The
northern group being the ancestral stock of the birds with streaked abdomens, and
the southern group of those with unstreaked abdomens. This split taking place
in an even wetter era than Stage 2 with a wide spread of forest and woodland over
most of central Africa. Proto-streptophorus still isolated south of the Sahara.
Stage 4. (East). A return of arid conditions with increased acacia allows the
northern streaked birds to spread down the coast, and the southern, unstreaked
birds to spread north up the Luangwa valley, through western Tanganyika and
Kenya and the Abyssinian Rift. The two types meet first in the north, and in
Somalia they mix and interbreed to a limited extent forming a hybrid zone but
with streaked birds always predominant near the coast. Subsequently they meet
in northern Tanganyika and later still in southern Nyasaland, but do not interbreed.
Stage 4. (North). In a dry era the acacia belt and proto-streptophorus move south
to the ridge of high ground from the Cameroons to Uganda and north-western
Kenya (Map 6A).
Stage 5. (East). An increase in humidity with a spread of woodland in central
Africa cuts off the northern unstreaked birds, proto-grantn subsp., from the
southern unstreaked birds, proto-zambesiaej sephaena subsp.
Stage 5. (North). An increase in humidity, coming first to the low ground of the
Shari basin and Bahr-el-Ghazal, north of the ridge, isolates proto-streptophorus
along the ridge. Increased humidity and competition with indigenous woodland
species drive it into refuges at the extremes of its range where more arid conditions
persist.
THE RED-WINGED GROUP (MAP 7)
(F. psilolaemus, F. shelleyi, F. africanus,1 F. levaillantoides , F. levaillantii, F.finschi]
RANGE AND CHARACTERS
This is a very homogeneous group which ranges over most of eastern and southern
Africa from Eritrea to the Cape and westwards to Angola. The various species
have a complex distribution but are largely allopatric though the ranges of three
overlap in the Transvaal. Members of the group are found in a wide variety of
habitat and at any altitude, and even individual species are found in varying habitats
in different parts of their range.
1 F. afer of Praed & Grant — see appendix.
THE FRANCOLINS, A STUDY IN SPECIATION 153
All birds of the group have " quail-type " plumage on the backs, with a lattice-
work pattern of irregular white or buff bars and shaft streaks on a basic colour of
mixed black, brown, grey and chestnut: all have the under wing-coverts and part
of the wing chestnut (including the South African form F. africanus, commonly
known as the Greywing, in which the chestnut is reduced) : all have a marked facial
pattern but this, and the pattern on the underparts, varies in the different forms:
all have black bills, usually with a yellowish base, and yellowish legs, the males
usually with a single spur. The sexes are alike.
SPECIES AND POTENTIAL SPECIES
The distribution and relationships of the various forms of this group would be
easier to discuss if no previous attempt had been made to designate species or
subspecies in the East African forms. I cannot agree with any of the classifications
made for this group which seem to place too great a reliance on minor morphological
characters, particularly the size of the bill, without regard to the ecology of the
forms of their likely evolution. The rearrangement I propose I believe to be more
realistic in these respects, though I appreciate that conventional nomenclature fails
to give more than an inadequate picture, and it is necessary as well to discuss the
relationships in some detail. To facilitate the recognition of the species as re-
arranged they will be discussed under separate headings and I have proposed new
common names for each.
F. psilolaemus — The Montane Redwing.
F. psilolaemus comprises the montane populations of Kenya, Uganda and
Abyssinia known by Praed & Grant as F. shelleyi theresae, F. s. elgonensis, F. afer
psilolaemus and F. a. ellenbecki.
All these birds are found in heath and grasslands above 8,000 ft. All have a high
proportion of rich chestnut in the wings and differ from other red-winged francolins
in having some barring on the tips of the primaries: the underparts are rich buff
mottled with chestnut and with some dark brown or black markings: the facial
pattern is broken and ill denned. Birds from the Shoa mountains round Addis
Ababa (psilolaemus) are small with small bills (wing ^164-174, bill ^30-34), and
have blackish spots on the throat, indistinct black spotting on the upper breast
and rather sparse chestnut and blackish patterning on the abdomen. Birds from
the Arussi plateau south of the Abyssinian Rift (ellenbecki) are slightly larger than
Psilolaemus, darker on the head and back, the lighter markings being greyer: below
the spots on the throat and breast are darker and the patterning is stronger and
richer in colour. Birds from Mount Kenya and the Aberdares (theresae) are large
(wing (£174-191, bill ^35-38) : they have only occasional speckles on the throat
but are heavily spotted on the upper breast : they are darker above than Abyssinian
birds and the chestnut markings on the abdomen are richer. Birds from Mount
Elgon (elgonensis) are similar to theresae but darker and richer.
F. shelleyi — Shelley's Redwing.
F. shelleyi comprises the forms known by previous authors as F. s. shelleyi,
ZOOL. 10, 2 II
154 B- P. HALL
F. s. whytei, F. afer uluensis, and F. a. macarthuri Van Someren, 1938, but excludes
theresae and elgonensis which were previously ascribed to shelleyi (see above).
All the birds included in this assemblage differ from F. psilolaemus in having the
breast maroon, chestnut and grey, unmarked with black below the necklace and
gorget : the remainder of the underparts are patterned all over with black and white,
the pattern basically of white spots on a black ground, but the spots frequently
merging to form irregular bars. In Kenya and northern Tanganyika birds of this
type with rather small bills, white throats, well-defined facial pattern and necklace
and rather small patterning on the abdomen are found on hillsides, grassy plateaux,
and clearings in montane forest from Mount Kenya southwards to the Crater
Highlands and Mount Meru. On Mount Kenya and the Aberdares they are found
at lower levels than F. psilolaemus keeping below the montane forest. Since they
are confined to uplands above 3,000 ft., their distribution is necessarily discontinuous
and some divergence is shown by isolated populations, birds of the Loita Plains in
south-western Kenya being rather greyer and those of the Chyulu Hills (macarthuri)
rather darker than birds from central Kenya (uluensis).
The most southern representative recorded of this small-billed form is from
Mbulu at 4,900 feet in the highlands south of Lake Manyara in northern Tanganyika.
Fifty miles south-east at Salanga on the Bereku Ridge at about 5,500 feet a large-
billed bird has been collected which has larger, more open, patterning on the
abdomen: between the two localities there is lower, rather open country probably
unsuitable to these francolins. Birds of the large-billed type are found on the
Uganda/Tanganyika borders and at scattered localities through western and
southern Tanganyika, Mozambique, southern Nyasaland, southern Northern
Rhodesia, Southern Rhodesia the Transvaal and Natal. The small-billed, small-
patterned form of Kenya (uluensis) and the large-billed, large patterned form of
Tanganyika (shelleyi) have been regarded as belonging to different species and the
proximity of the two in the Mbulu area lends weight to this view. Nevertheless,
they seem to have similar ecological requirements and field habits; and, while in
the Mbulu area the topography of the country may inhibit interbreeding, there is a
bird from Amani, 200 miles to the west, which is intermediate between the two
forms in the patterning on the abdomen, though closer to shelleyi in bill size, which
suggests there may be no intrinsic barrier.
The long-billed shelleyi is found in varying habitats throughout its range. In
southern Nyasaland, as in northern Tanganyika, it may be found on montane grass-
land up to 7,000 feet, but it is also found in Nyasaland and Northern Rhodesia in
grass under brachystegia or even mopane at low altitudes, though it is nowhere
common. In Southern Rhodesia it is common in Mashonaland and the Midlands,
in open grass of the plateau, but scarcer westwards in Matabeleland (Smithers
et al, 1957: 46). It occurs at least as far south as Pretoria and Legogot, in the
Barberton district of the Transvaal, in presumably the same type of country as in
Southern Rhodesia, but in Natal is found in acacia country below 2,000 ft. along
the foothills of the Drakensburg.
In northern Nyasaland, the south-eastern Congo and northern Northern Rhodesia
the species (as proposed) is represented by another long-billed form (like shelleyi), but
THE FRANCOLINS, A STUDY IN SPECIATION 155
with a small patterning (like uluensis) and which differs from both in having a buff
throat with the facial pattern and necklace freckled and indistinct. This form
(whytei], like shelleyi, is found at varying altitudes and in both brachystegia and
montane grassland and, though there is no ecological barrier between them and
they have been collected from within 40 miles of each other at Lundazi and Mzimba,
no intermediates are known (Benson, 1951: 79). Benson tells me that he believes
this is more likely to be due to lack of collecting and to the scarcity of both forms,
rather than to the existence of any barrier to interbreeding. The situation may be
found to be comparable to that between uluensis and shelleyi in northern Tanganyika
but further research is required.
F. africanus (F. afer) — the Greywing.
I regard the South African Greywing as a monotypic species, and do not associate
any of the northern short-billed forms with it. It differs from F. shelleyi in having
a white throat flecked with black, the facial pattern obscured by black-and-white
freckling, the patterning on the underparts very small, and the rufous in the wing
very much reduced. The bill is small as in uluensis. It is found in the grasslands
of the mountains of the southern Transvaal, Orange Free State and Cape Province,
reaching Natal on the high spurs of the Drakensburg. McLachland & Liversidge
(1957 : 94) show an overlap in the ranges of F. shelleyi and F. africanus in the
southern Transvaal but I cannot substantiate this, the most northerly specimens o'f
africanus recorded being from Potchefstrom and Wakkerstrom, 100 miles south of
shelleyi at Pretoria, and in rather different country.
In Natal Vincent tells me there is a clear altitudinal distinction between them,
africanus being found above 6,000 ft. and shelleyi below 2,000 ft., while the inter-
mediate levels are occupied by a third member of the group, F. levaillantii. Further-
more F. shelleyi has an earlier breeding season than the other two species, which
would contribute to segregation.
It will be appreciated therefore that africanus has, in many respects, the same
relationship to shelleyi in the Transvaal as shelleyi has to whytei in Northern Rhodesia
and to uluensis in Tanganyika, in that it appears to be completely allopatric and to
exhibit distinct morphological differences. But, whereas there appear to be no
ecological barriers between shelleyi /uluensis and shelleyi /whytei and the morpho-
logical differences are slight, there is a distinct ecological barrier between shelleyi
and africanus and morphological differences are greater, representing a greater
degree of divergence. This to me is just sufficient to warrant giving africanus
specific rank.
F. levaillantoidss (olim F. gariepensis) — Acacia Redwing.
F. levaillantoides comprises all the subspecies of southern Africa usually ascribed
to this species and in addition the northern populations known by Praed and Grant
as F. afer stantoni, F. a. friedmanni, F. a. archeri, F. a. lorti and F. a. gutturalis.
It will be seen from the foregoing that I regard the red-winged francolins of the
northern and southern acacia steppe as conspecific. They differ from F. psilolaemus
and from F. shelleyi in being paler and less patterned below, lacking on the abdomen
156 B. P. HALL
the rich chestnut markings of psilolaemus or the black-and-white patterning of
F. shelleyi, having instead sparse chestnut and blackish flecks or streaks, more
common in the northern birds.
In southern Africa they are found in the arid zones from coastal Angola through
South West Africa, Bechuanaland, Orange Free State and the Transvaal to the
east coast in extreme southern Mozambique, associated, at varying altitudes, with
open grassland, hillsides and kopjes, and edges to limestone pans. There are
sight records (not included on Map 7) from the Lower Orange River (Winterbottom
& Courtenay-Latimer, 1961: 11-12 and in litt.}. The range of levaillantoides
overlaps that of africanus in the Orange Free State and that of shelleyi in the
Transvaal. There is little information on their relationship here (perhaps because
their numbers have been heavily reduced since the country was settled) but it can
be expected that levaillantoides would be found at lower altitudes than africanus,
and in drier areas than shelleyi.
In the northern acacia belt the red-winged francolins are found in much the
same habitat as southern birds, in sparse grass on rocky hillsides and mountains
between about 2,000 and 7,000 feet. In central Abyssinia, where the Acacia
Redwing is found in the same areas as the Montane Redwing, it occupies the lower
slopes and F. psilolaemus the higher (Friedmann, 1930: 112-116).
In northern Abyssinia and Eritrea there is an isolated population (gutturalis)
found in scrub above 6,000 feet (Smith, 1957: 20). These birds appear to be a
link between the acacia species and the Montane Redwing F. psilolaemus, for, like
the montane bird, they have the facial pattern freckled and ill denned. They also
differ from other Acacia Redwings in having the abdomen regularly marked with
heavy blackish streaks on the feather centres, though the actual shafts are white,
while the flanks are broadly streaked with chestnut. However, in colour and in
the distribution of patterning below they seem closer to the Acacia Redwings and
are tentatively regarded as conspecific with them.
Otherwise variation in pattern among the Acacia Redwings is slight. All have
the facial pattern and necklace well defined, but in birds from southern Angola
(jugularis) south at least to the Cunene, the black-and-white necklace is enlarged
on the breast to a broad gorget. The birds from the Benguela area are exceptionally
small, and also rather grey, while Cunene birds are larger and rufous. Throughout
both the northern block and other members of the southern blocks there is similar,
very marked variation in colour between grey and rufous populations, largely
associated with ecology, but there is remarkably little consistent difference between
northern and southern birds in spite of the great distance separating them; those
from the north have commonly, however, fine dark markings on the abdomen that
are usually lacking in those from the south and the abdomens are whiter, less buff.
The greyest populations are found in the mountains of eastern (British) Somaliland
(lorti), in the Kalahari (kalaharica — see appendix), and in the Benguela district
(as already noted) . The most rufous birds are found in the Transvaal and Orange
Free State (levaillantoides}, while in both South West Africa (pallidior} and southern
Abyssinia and southern Sudan (archeri) both rufous and greyish micro-populations
are found haphazard (see appendix).
THE FRANCOLINS, A STUDY IN SPECIATION 157
It will be seen that, as far as the northern populations of the red-winged francolins
are concerned, the subspecies of the acacia zone could be considered conspecific
with F. shelleyi, since they are aUopatric. However, since in southern Africa
F. levaillantoides and F. shelleyi both occur in the Transvaal, two species must be
recognised, and as the northern acacia populations are closer both morphologically
and ecologically to the southern acacia birds than to F. shelleyi, it seems logical to
associate them in the same species.
F. levaillantii — Levaillant's Redwing.
I have made no rearrangement in this species, which is universally recognised
as comprising scattered populations from north-western Kenya to the Cape and west-
ern Angola and which differs from all other red- winged francolins in having an ochre
collar and the sides of face and edges of the throat ochre inside the black-and-white
facial pattern. Such populations are found in north-western Kenya, where west
of the Rift F. levaillantii replaces F. shelleyi in grasslands and valleys of the plateaux
between 6,000 and 8,000 feet. In Uganda and Ruanda it is found at varying
altitudes being common in the lower grasslands west of Lake Victoria (south of
Bukobo) and on the plateaux westwards to Lakes Edward and Kivu.
It is absent from most of Tanganyika but reappears on the Njombe and Iringa
mountains in the south-west, and on the Nyika Plateau of northern Nyasaland,
where it is common above 7,500 feet alongside the rarer F. shelleyi whytei (Benson,
in litt.}. Another isolated population is found on the Kibera Plateau in Upemba,
southern Congo, at about 6,000 feet (Verheyen, 1953 : 256), and scattered populations
are found in the grasslands of the Balovale and Mongu districts of north-western
Northern Rhodesia westwards through Angola to the Mombolo plateau and the
Bailundu highlands. In southern Africa it is found only from the Zoutspansberg
southwards through the eastern Transvaal, Natal (between 2,000 and 6,000 feet),
Basutoland and the coastal districts of eastern Cape Province, keeping to the
moister grasslands and even reed-beds in river valleys. It seems, therefore, in the
Transvaal, Natal and Orange Free State, where the four species of Redwing occur,
that F. levaillantii is associated with the wettest habitat.
The distinctive ochre patterning of the face and throat of F. levaillantii has
already been noted : it is otherwise most like F. levaillantoides but is long-billed
and darker and richer in colour, with the black-and-white patterning of the necklace
and gorget extending in a complete collar round the hind neck below the ochre
collar, and the black-and-white stripes from above the eye, which in levaillantoides
and shelleyi run down the side of the face, in levaillantii run behind the head to join
at the back. In spite of the wide range and broken distribution there is little
variation between the populations but in the populations of southern Africa and of
the Nyika plateau these stripes continue down the back of the neck to join the
black-and-white collar, while in the western and northern populations the ochre
collar is unbroken. The Nyika birds (crawshayi) are richer in colour than those of
southern Africa (levaillantii), with more rufous in the wing, and with more black
markings on the abdomen, while the northern birds without the stripe on the hind
neck (kikuyuensis) are otherwise similar to crawshayi.
I58 B. P. HALL
F. finschi — Finsch's Redwing.
The last of the red-winged species, F. finschi, is monotypic. It has a very limited
distribution being recorded only from about six localities in western Angola, in a
strip about 200 miles long from near Nova Lisboa in the south to Vila Salazar in
the north, and again from Brazzaville, in the Congo, 350 miles to the north. On
Mount Moco in the Bailundu highlands it was found in the brachystegia woodlands
and on the bare upper slopes between 6,500 and 7,000 feet (Hall, i96oa: 408) but
the specimen from Vila SaJazar (American Museum of Natural History) must have
been obtained in very different habitat at not more than 2,000 feet, and at Brazza-
ville Malbrandt and Maclatchy (1949: 142) found it in grassland with neighbouring
woods in which the birds took refuge.
It is possible that the different habitats of F. finschi may be partly due to com-
petition with F. levaillantii, which occupies grasslands in the southern part of the
range, but is not found north of the Mombolo plateau.
F. finschi differs from other red-winged f rancolins in lacking any black-and-white
patterning on the face and neck and having the breast grey: it is closest to F.
levaillantii in having a long bill and the sides of the face and border of the throat
ochre, but in F. finschi the ochre is not so extensive on the hind neck : the abdomen
is a mixture of buff and pale chestnut.
OTHER VARIATION
Considerable individual variation is to be expected in any birds as intricately patterned as
the red-winged francolins and is found in all species. There is some local variation, usually
in an increase or reduction of rufous and grey in the plumage, associated with ecology. This,
as noted, is particularly marked in F. levaillantoides, but since the problems it raises are largely
nomenclatorial it is discussed fully in Appendix 2.
EVOLUTION
The complex pattern displayed by the seven species of the Red-winged Group is
not easy to interpret but I believe may be partly explained by comparison with a
rainfall map (Map 7A) . From this it will be seen that the range of F. levaillantoides
falls largely in the parts where the rainfall is below 20 inches in the year, the range
of F. shelleyi falls between the 20-40 inch isohyets, the range of F. levaillantii falls
between the 40-60 inch isohyets, while the range of F. finschi coincides remarkably
with the only two patches south of the Congo rainbelt where there is over 60 inches
in the year. Thus a slight increase or decrease of rainfall may be expected to favour
the spread of one species at the expense of others. The discontinuous distribution
of all species, and the scarcity of red-winged francolins in so many parts of their
range, suggests that the present cannot be regarded as a favourable era for any
members. This might be due to competition with other groups, but if so it could be
expected that F. levaillantoides would be more common in the Kalahari, where
there is no competition. From the association of so many members of the group
with mountains I believe that a colder era would be more favourable to the group
as a whole, and it was in some such era or eras, with fluctuations in rainfall, that
THE FRANCOLINS, A STUDY IN SPECIATION 159
continuity in the ranges of the various species was achieved. Conversely warmer
eras could be regarded as least favourable and ones most likely to produce isolated
populations in which speciation developed.
Even with this hypothesis as a basis a good many interpretations of the pattern
present themselves which are largely dependent on which species one believes are
most closely related. The following sequences are put forward tentatively on the
premise that plumage pattern is the best guide to relationship. I can offer no
explanation of the variation in bill size in terms of evolution and relationship but it
is worth noting that in this group it is the species and, in shelleyi, the subspecies,
associated with the higher rainfall areas that have the long bills. The length may
perhaps be correlated with the hardness of the ground in which the birds search for
food.
The distinctive patterning on the abdomens of F. africanus and F. shelleyi suggests
that their common ancestor split at an early date from the rest of the group. The
present distribution suggests they developed in the south ; later spreading north to
Kenya in a favourable era; later splitting into two blocks, proto-africanus and
proto-shelleyi ; later proto-shelleyi splitting into three from which the subspecies
shelleyi, whytei and uluensis developed. A favourable era must have followed in
which their distribution was continuous over the present range.
The species with relatively unpatterned abdomens may have originated in the
north. A warm era could be expected to drive members of this block to the
extreme north of the range, and also to high altitudes: it can be postulated that
proto-psilolaemus developed in such an era to specific status and became adapted to
montane conditions : later in a favourable era other members of the northern block
spread south and west as far as Angola and the lower Congo. A subsequent un-
favourable era isolated first proto-finschi in the Angola mountains, then proto-
levaillantii in the mountains of central Africa, and proto-levaillantoides in Abyssinia,
Somaliland and Eritrea: in isolation each became adapted to different degrees of
humidity: subsequent fluctuations in rainfall allow first proto-levaillantoides to
spread widely through the continent and then proto-levaillantii. The small degree
of divergence shown between the isolated populations of both species (with the
exceptions of F. levaillantoides gutturalis in Eritrea which perhaps was isolated
earliest) suggests that these dry and wet cold spells were comparatively recent.
A summary of the evolution postulated is as follows: —
Stage i. The group widespread in eastern Africa in a cold era.
Stage 2. The group split into a northern and southern block.
Stage 3. The southern block spreads north, at least to Tanganyika, and the northern
block south to Kenya (Map 76).
Stage 4. Warm and unfavourable, both blocks split into two : in the south proto-
africanus isolated in the mountains of South Africa, proto-shelleyi possibly in the
mountains of Nyasaland: in the north proto-psilolaemus isolated at high altitude,
possibly on Mount Kenya or Elgon, and the ancestral stock of the other species
isolated in the extreme north.
Stage 5. Favourable. Proto-shelleyi spreads, possibly to the limits of its present
160 B. P. HALL
range. Members of the northern blocks (with unpatterned abdomens) spread south
and west to Angola.
Staged. Unfavourable. Proto-finschi isolated first in Angola: proto-levaillantii
isolated next in some mountains of central Africa not occupied by proto-shelleyi,
while proto-levaillantoides is cut off in the north. Each becomes adapted to different
requirements of humidity.
Stage 7. In a long cold era favourable to the group as a whole, fluctuations in rainfall
and temperature favour each species in turn. In some unusually cold spell proto-
psilolaemus spreads to cover a large part of Abyssinia and Kenya but is later split
into isolated populations in the highest mountains. Proto-shelleyi following a
favourable era is split into three, from which the subspecies shelleyi, whyteiand
uluensis develop almost to specific status. An unfavourable spell in the north
isolates proto-gutturalis in Eritrea from the main body of proto-levaillantoides: in
a subsequent dry spell (during which the acacia spreads to link the northern and
southern belts) proto-levaillantoides spreads south to Bechuanaland. In a wet
spell pToto-levaillantii spreads at the expense of shelleyi and levaillantoides, and
proto-finschi spreads: where the two come together finschi is pushed into the
mountains.
Stage 8. The present. Too warm to be favourable to any members of the group,
the ranges of all species being discontinuous and the birds uncommon in most
areas.
THE RED-TAILED GROUP (MAP 8)
(F. coqui, F. albogularis, F. schlegelii}
RANGE AND CHARACTERS
This is a group of small francolins (wing under 150 mm.) which ranges from
Senegal to the Sudan, and from central Kenya west to the central Congo and
Angola, and south to the Transvaal and Natal: there is an isolated population in
the Rift of Abyssinia. Members of the group are found in woodland, savanna and
steppe and, in parts of the range, often on the same ground as other francolins, which
suggests that their smaller size may make them non-competitive. The group is
formed of three species which are allopatric except in the southern Congo and
eastern Angola where isolated populations of albogularis are found within the range
of coqui.
All members of the group have a considerable amount of ochre on the sides of the
face, spreading as an ill-defined collar on the hind neck : the crown is plain brown or
chestnut, contrasting with the back which has a quail-type pattern (except in the
females of schlegelii in which the striations and cross bars are reduced to a minimum),
varying in colour from grey and black to vinous rufous: in the redder birds the
rufous in the tail is conspicuous but in some of the greyer forms is no more than a
pinkish wash on the outer feathers: the underparts are variable but all birds have
a white or buff throat contrasting with the breast: all have black bills with some
yellow at the base and yellow legs : the males have a single spur.
THE FRANCOLINS, STUDY IN SPECIATION r6r
SPECIES AND POTENTIAL SPECIES
F. coqui is the most widespread of the three species which comprise the group.
It is found in short grass in brachystegia woodland from southern Uganda and
northern Tanganyika, southwards to the Rhodesias and westwards to Brazzaville
and the edge of the Angola escarpment, but is absent from mountains and from the
drier acacia steppe of Tanganyika. South of the brachystegia belt it is found in
the acacia savanna skirting the Kalahari, from northern South West Africa to the
Transvaal, and in the coastal districts of Natal. Scattered populations in Kenya,
southern Abyssinia and the Harar area are also found in acacia (Benson, 1945 : 390)
finding it common around Yavello and Mega in " open short-grassed country with
scattered low thorn bushes at 4,000-4,500 feet ".
In West Africa there is a population in the acacia belt of Senegal, the Upper
Volta and northern Nigeria that is isolated by 1,000 miles (and the Lower Guinea
forest) from the Brazzaville birds, and by nearly 2,000 miles from the Abyssinian
birds. In this area coqui is found commonly in cultivations, often alongside F.
bicalcaratus (Blasdale: unpublished). However, it seems that at Potiskum, 150 miles
east of Kano in northern Nigeria, coqui is replaced abruptly by F. clappertoni which
occupies the acacia belt eastwards to Abyssinia (Hollis: unpublished). In
Abyssinia clappertoni occupies the northern and western areas to the centre of the
Rift but is replaced by coqui in the Harar area in the east and in the Mega/Yavello
area of the south.
In coqui the quail-type patterning on the back is very consistent and well defined,
though the basic colour, and the colour of the head, vary from grey to brown or
rufous in different populations. The sexes differ, the males having the sides of the
face and throat ochre or light buff without a black eye-stripe or necklace, the
females having the sides of the face similar in colour to the males but the throat
whiter and having a black necklace and a black eye-stripe that continues as a black
line down the sides of the face. The males also have the breast barred while in
the females the barring is replaced or overlaid by a pink or grey wash.
There is little significant geographical variation in birds of the southern part of
the range, from Angola, the Congo, central Tanganyika and coastal Kenya south to
South Africa (coqui}. All have grey wings and barred abdomens (though the degree
of barring is variable) and females have a pink wash on the breast. In the
Abyssinian population (maharao) the wings are rufous and both sexes have light
narrow barring from breast to abdomen, the females having only a faint wash of
pinkish grey over the barring on the breast. The isolated West African population
(spinetorum) also has rufous wings, but is pale above, washed with pink, and there
is no barring on the abdomen in either sex: the breast of male is broadly barred
and that of the female plain and pinkish-grey. In Uganda, Kenya and Tanganyika
the pattern of variation is confused. East of the Rift in Kenya, south to the Arusha
area of Tanganyika, there is a population (thikae, Grant & Praed, 1934) known only
from a few specimens, which shows some approach to the Abyssinian birds in having
rufous wings, light barring, and vestigial barring on the breasts of the females, but
with no barring on the abdomens in either sex. From the Rift westwards to the
eastern shores of Lake Victoria the birds (hubbardi) are similar to both the West
162 B. P. HALL
African spinetorum and the eastern thikae in having plain abdomens but the wings
are grey like the southern coqui, the breasts of the females are plain and grey, and the
whole upperparts are rather dark and grey. Birds from west of Lake Victoria
(ruahdae) are close to the southern coqui, but have the black barring broader and more
widely spaced. These birds intergrade with hubbardi in the Tabora area.
The second species in the group, F. albogularis, has a scattered distribution.
There is a population in the Gambia, found mostly on the edges of cultivation, but
it is rare (Hopkinson, 1923: 129). Other populations are found on the coastal
plain near Accra and in the hinterland of Ghana, eastwards through Nigeria to
Garoua in the (French) Cameroons. Field notes on these birds are scarce and
somewhat contradictory, Serle (1957:389) finding them common in "the open
savannas and rolling hill country " of Eastern Nigeria, whereas Blasdale tells me
that further north he has found them only in thick woodland where there is
abundant Isoberlinia doka. Most localities from which they have been collected
lie within the woodland belt (Map n, Type 17) but Garoua lies within the acacia
savanna belt (Map n, 20) and the specimen was obtained in " open, sandy, sparsely-
treed country " (Good, 1952: 67).
F. albogularis is absent from the woodland and savanna belt north of the Congo
forest (where F. schlegelii is found) but reappears again 1,500 miles away in the
south-eastern Congo on the plateaux of Marungu and Kibara, in the Upemba National
Park, on the plains of the Balovale district of Northern Rhodesia, and on the treeless
dambos of the Zambezi/Congo watershed in eastern Angola.
Above F. albogularis is very like F. coqui but the quail-type patterning is less
well defined in the females, the shaft streaks and barring being narrower: the wings
are more rufous (similar in this respect to the east Kenya population, thikae). As
in coqui the females have a black facial pattern and necklace, but it is ill defined,
and the males have none, though they differ from coqui in having a white throat
contrasting with the ochre cheeks, as in the females. In the rest of the underparts
the males of albogularis are quite distinct from any form of coqui being chestnut on
the breast with ochre shaft streaks and rich ochre on the abdomen, lacking any dark
barring. The females on the other hand are barred, in greater or lesser degree
according to the population, with a faint wash of pink or rufous on the breast, and
resemble closely the females of the Abyssinian and east Kenya populations of
coqui. The population of albogularis in which the females are least patterned is
in the Gambia (nominate albogularis) the barring being light and restricted to the
breast and flanks (though varying individually). In birds from Ghana to the
Cameroons (buckleyi) the barring is generally heavier, usually with only the centre
of the abdomen unpatterned. Birds of both sexes from the populations south of
the Congo forest are slightly larger and more richly coloured than those from West
Africa and in the females of the south-east Congo (dewittei) the barring is heavier
but no more extensive than in buckleyi: the darkest birds with the most heavily
barred females are found in eastern Angola and possibly the Balovale district
(meinertzhageni White, 1944 — see appendix).
The third species of the group, F. schlegelii, has been considered by Chapin and
Peters to be a subspecies of F. coqui. I believe it is distinct enough to warrant
THE FRANCOLINS, A STUDY IN SPECIATION 163
specific status and furthermore is closer in several respects to F. albogularis.
It is a rare bird, found scattered through the woodland and savanna belt from the
eastern Cameroons to the western Bahr-el-Ghazal. From what Cave tells me it
seems to be more consistently a woodland bird than albogularis and he has the
impression that it is found in rather denser woodland than the sympatric F.
icterorhynchus of the Vermiculated Group, especially where there is Isoberlinia doka.
Above schlegelii is closer to albogularis than coqui, though the quail type patterning
is much reduced and the sexual dimorphism is more pronounced, the males having
broad white shaft streaks but few transverse bars while the females are almost
unpatterned: both sexes have a soft vinous wash. Below the male of schlegelii is
like that of southern races of 'coqui, with ochre sides of the face and throat and
narrow black and white barring over the breast and abdomen : the female resembles
the male on the head and throat but has the breast grey, patterned with cream
triangular marks at the base of the feathers, the flanks sparsely barred, and the
centre of the abdomen plain cream.
OTHER VARIATION
In the southern populations of F. coqui there is local and ecological variation in size and
colour; exceptionally small birds are found in coastal Kenya and Natal; grey, pale birds in
Bechuanaland (vernayi) and pale, rather pinkish birds in South West Africa (hoeschianus
Stresemann, 1937) : other exceptionally grey birds occur in southern Mozambique (Coguno) and
may be expected locally in dry conditions, while pockets of more richly coloured birds are
associated with richer vegetation. There is otherwise little variation between birds from Natal
and the Transvaal north to coastal Kenya and west to Northern Rhodesia, except in the
extreme west: here and in Angola the birds are consistently less rufous and the bars on the
underparts are narrower and rather closer together (angolensis) . Birds from Kasai and
western Katanga (kasaicus White, 1945) are similar to coqui above and angolensis below: they
intergrade with coqui in the eastern Katanga.
In F. albogularis of West Africa individual variation in patterning has already been noted.
Serle (1957: 388) also found the general colour individually variable between rufous and grey,
though there seems a tendency for the coastal population of Accra to be greyer than inland
birds.
Individual variation is also found in F. schlegelii especially in the females (see Tray lor, ig6ob:
86).
EVOLUTION
The most striking features of the pattern presented by the Red-tailed Group
are the enormous distances separating the north-western and south-eastern sub-
species of both F. albogularis and F. coqui: also, in F. coqui, the complexity of
variation in East Africa in contrast to the uniformity of southern birds. I feel
confident that the discontinuous range of F. albogularis is largely attributable to
the presence of F. schlegelii in the intervening segment of the same vegetational belt,
and suggest that the ancestral stock of the two species once had a continuous
distribution in the savanna and woodland encircling the Congo forest. To account
for the split into two species I suggest this era was followed by a very dry period
in which the forest was divided into a western and eastern block each encircled by
woodland and separated by a corridor of acacia linking the northern and southern
164 B. P. HALL
acacia belts. It can reasonably be postulated that proto-albogularis developed in
the belt round the western forest and proto-schlegelii in the belt round the eastern
forest (Map SB). When the two forest blocks rejoined with a return of humid
conditions and the surrounding belts became one again, the central segment of the
belt was occupied by proto-schlegelii and the western and southern segments by
proto-albogularis.
It is more difficult to interpret the pattern of F. coqui but a logical sequence can
be found which will fit into that postulated for the other species. Proto-coqui
may have been separated from the ancestral stock of the other two species at a
time of spread of montane forest and developed in the east while proto-albogularis /
schlegelii developed in the north. With the retreat of the forest proto-coqui spread
south and west chiefly in acacia savanna. In an unfavourable spell, either too
wet or too dry, proto-maharao in Abyssinia was isolated. In the very dry era
postulated above, in which there was a corridor of acacia in the central Congo,
proto-coqui could have spread into the northern acacia savanna belt. Subsequently
with a return of humid conditions the northern population would be cut off from
the southern by woodland and forest. The southern populations nearest the Equator
prove adaptable so that, with the return of woodland and savanna and the in-
digenous albogularis to the southern Congo and Angola, coqui competes successfully,
occupying the woodlands and albogularis the more open grasslands and plateaux.
Meanwhile the isolated northern population diverges from southern birds, develop-
ing the distinctive unpatterned abdomen in both sexes, and spreads through the
northern acacia belt west to Senegal and east through the Sudan to Kenya and
Uganda. Here it encounters both the Abyssinian maharao and the southern coqui.
The characters of the present populations of Kenya (hubbardi and thikae) and of
southern Uganda (ruahdae) suggest that some interbreeding took place at some
period.
The present appears to be favourable to the Vermiculated Group in the north,
at the expense of the Red-tailed Group. In the immediate past it seems that
F. clappertoni has spread west through the acacia belt in the Sudan, eliminating
coqui between the Nile Valley and northern Nigeria. In the woodland and savanna
belt F. bicalcaratus flourishes in the west and F. icterorhynchus in the east, and
competition with these two may account for the scarcity and discontinuous dis-
tribution of albogularis and schlegelii. However, in West Africa coqui, though
apparently unable to compete with clappertoni, is yet able to live on the same
ground as bicalcaratus on the southern fringe of the acacia belt. The populations of
coqui in eastern and southern Africa also seem able to co-exist on the same ground
as other francolins.
A summary of the evolution postulated is as follows : —
Stage i. The group develops in savanna, perhaps in north-eastern Africa.
Stage 2. An unfavourable era, possibly a spread of montane forest. The group
splits into two with proto-coqui isolated in the east and proto-albogularis I schlegelii
to the north of the forest.
Stage 3. More favourable. Proto-coqui spreads south and west in rather dry
THE FRANCOLINS, A STUDY IN SPECIATION 165
savanna, mostly acacia. PToto-albogularis/schlegelii spreads in the wetter savanna
and woodland encircling the lowland forest of the Congo Basin (Map 8 A).
Stage 4. A very dry era. The Congo forest is divided into a western and eastern
block, each surrounded by its own woodland and savanna belt, with a corridor of
acacia between. Proto-albogularis develops in the belt surrounding the western
block of forest, proto-schlegelii in the belt surrounding the eastern block. Proto-
coqui spreads through the acacia corridor into the northern acacia belt. In an
unfavourable period in the north-east proto-maharao becomes isolated (Map SB).
Stage 5. More humid. The two blocks of forest rejoin and the two savanna belts
become one again, the eastern segment being occupied by proto-schlegelii and the
western and southern segments by proto-albogularis. In the southern segment, as
woodland and savanna return to districts previously acacia, proto-coqui is not
pushed out but becomes adapted to woodland and competes successfully with
proto-albogularis which takes to the more open grasslands. In the north proto-
spinetorum subsp. spreads west to Senegal and east to Abyssinia and Kenya and
diverges from the southern birds. There is limited interbreeding where the northern
birds re-encounter other members of the species in the east.
Stage 6. Favourable for the Vermiculated Group, especially in the north. F.
clappertoni spreads, eliminating F. coqui from the acacia belt in the Nile Valley and
Sudan westwards to northern Nigeria. In the woodland and savanna belt F.
bicalcaratus and F. icterorhynchus flourish at the expense of F. albogularis and
F. schlegelii. In the south F. coqui co-exists more successfully with francolins of
other groups.
SPECIES NOT INCLUDED IN ANY GROUP (MAPS 9 & 10)
Francolinus lathami
F. lathami is a forest francolin ranging from Sierra Leone to western Uganda and
the southern Sudan.
It appears at first sight quite unlike any other African francolin (closer, except in
size, to F.francolinus of Asia), being, in the male, largely black below with regular
white spots, in the female brown with irregular white spots, the throat black and
face patterned, the upper parts mottled rufous and brown with some white streaks
on the mantle, the bill black, finer than in other francolins. However, comparison
of a pair of lathami with a pair of schlegelii suggests that lathami may be more
closely related to the Red-tailed Group than to any other francolin. It will be
seen that the pattern and the colouring on the upper parts are basically the same,
though lathami, as would be expected in a forest bird, is darker and more mottled.
Below the patterning of the females has some similarity in the irregular, rather
triangular shaped spots: while the spotted male of lathami looks very different
from the black-and-white barred male of schlegelii, this may not be a very significant
difference for in odd abdomen feathers of several lathami the spots are less regular
and merge as vestigial bars. The chief differences in fact lie in the shape of the
bill and the pattern of the face and throat. Furthermore the first plumage of
lathami chicks is very similar to the plain vinous mantle of the female schlegelii.
166 B. P. HALL
While being confident that there are affinities between lathami and schlegelii I
am not prepared to guess at what period in history proto-lathami became isolated
and adapted to forest. The likeness to one particular species in the Red-tailed
Group suggests that its isolation could have been subsequent to the emergence of
species within the group, the adaptation to such different habitat accelerating diver-
gence. On the other hand F. schlegelii is the species of the Red-tailed Group that
lives in the thickest woodlands and, if proto-lathami had broken from the ancestral
stock of the Red-tailed Group before the three species had diverged, the similarities
between lathami and schlegelii might be due to convergence. For this reason, and
with regard to both its morphological and ecological distinctions, I feel it best to
treat lathami independently.
It will be seen that I am not prepared to regard the black-and-white spotting
in the males of lathami as indicative of relationship with F.francolinus. It has
been amply demonstrated in the discussion on groups that the pattern of the
underparts is of less significance than size and the pattern above, and while it is
interesting that such a striking pattern is repeated in the genus in two such different
birds, it cannot be relevant to the grouping of species.
From the distribution plotted on Map 9 it would appear that there is a wide gap
between the populations of the eastern and western Congo, but this is more likely
to be due to lack of collecting in the central area than to a genuine break. There
is little geographical variation, in the males of eastern birds (schubotzi) the black-
and-white pattern extends further down the abdomen, and the females have the
cheeks more rufous than grey, but there is considerable individual variation.
Francolinus nahani
This little francolin is confined to the heaviest forests of the north-eastern Congo
and Uganda. It is a distinctive bird, predominantly dark brown or blackish, with
white streaks on the underparts and some white spots on the wing-coverts and
flanks. The bill is black at the tip and crimson at the base; there is a patch of
crimson bare skin round the eye; the legs are crimson, and without spurs in either
sex.
It is difficult to assess its affinities with the other francolins. Superficially it
appears closest to F. lathami in its small size, general dark coloration and suggestion
of spotting, and they occupy the same type of habitat. On the other hand the
similarities are in those characters that are to be expected in forest birds, and could
well be due to convergence, and the fact that F. nahani in the whole of its range is
sympatric with F. lathami, without apparently being ecologically segregated, does
not suggest close relationship. I would suggest very tentatively that F. nahani
may have broken off at a remote period from the ancestral stock of the Scaly Group
(in the same way as I have suggested that lathami may have broken off from the
Red-tailed Group), and become adapted to heavy forest instead of the forest clearings.
I suggest this because nahani, although much smaller, seems to have some characters
in common with the Scaly Group as a whole and with each of the three individual
species. In both nahani and all species of the group the sexes are alike, and the
legs are red: nahani is like F. squamatus above, having an unpatterned head and
THE FRANCOLINS, A STUDY IN SPECIATION 167
vermiculated back; it is like F. ahantensis in the white streaking below; and like
F. griseostriatus in the crimson base to the bill. Furthermore in all species of the
Scaly Group there is a small area of bare skin below and behind the eye, though this
is not brightly coloured and usually overlooked.
Chapin considered the absence of spurs to be a generic character and proposed
the monotypic genus Acentrortyx for F. nahani, but the fact that F. pictus in India
is also without spurs, but yet is so closely related to F. francolinus as to hybridise,
demonstrates that this character is not necessarily significant. I see no other reason
for regarding F. nahani as generically distinct.
F. pondicerianus
F. pondicerianus is the smallest of the Asiatic Francolins and is widespread from
the shores of the Gulf of Oman eastwards through the plains of India and the dry
north-western districts of Ceylon, living in grassland or scrub. It appears to have
some affinities with both the Red-tailed and the Striated Groups of Africa and in
colour and pattern is closest to F. coqui, in particular to the geographically nearest
form F. c. maharao of Abyssinia. It has " quail-type " plumage on the back,
but is duller than coqui, with broader transverse white bars and narrower shaft
streaks, and with chestnut markings on a grey or huffish ground: the underparts
are whitish, irregularly barred with very fine blackish bars: the throat is buff
surrounded by a blackish necklace, which is smaller than that found in the females
of coqui and starts from the chin and not the sides of the bill. In other respects
it is closer to the Striated Group (F. sephaena and F. streptophorus) than to the
Red-tailed Group, having a longer and more graduated tail (the tail more than half
the length of the wing), and stronger (though not necessarily longer) legs and larger
feet, the legs being reddish as in F. sephaena. The bill is distinct from either group
being brownish in colour and heavier in front of the nostrils. The tail in colour
combines characters of both groups being reddish at the base but otherwise largely
blackish.
The affinities with two of the African Groups suggest that the ancestral stock
of F. pondicerianus was isolated from the African stock before the Red-tailed and
Striated Groups had diverged and for this reason I have discussed it independently.
It seems that, while the African stock underwent many vicissitudes of climate with
consequent isolating and regrouping of populations resulting in speciation, the
Indian stock had a comparatively tranquil history for in pondicerianus there is
little variation, and that mostly clinal and ecological: birds from the dry districts
of Persia and Baluchistan, east to the western Punjab (mecranensis) , are the greyest
and palest, and those from southern India (pondicerianus) the darkest and least
grey with the greatest amount of chestnut markings. Those from northern and
central India.(interpositus) are intermediate. Ceylon birds (ceylonensis Whistler, 1941)
are greyer than either pondicerianus or interpositus but darker than mecranensis.
F. gularis
F. gularis is the swamp francolin of India, being confined to reeds and marshes
in the plains of the Ganges, Brahmaputra and their tributaries from central Uttar
168 B. P. HALL
Pradesh eastwards into Assam. It is a large francolin, comparable in size and
general proportions to the largest member of F. francolinus, but with rather a shorter
bill and larger feet (as would be expected in a marsh bird). It is, however, quite
different in colour and pattern being predominantly a brown and white bird, with a
barred back, streaked underparts and a rufous throat, wings and tail. The sexes
are alike.
Proto-gularis seems likely to have become isolated from other francolins in a dry
period and taken refuge in river valleys, becoming adapted to these conditions.
The dry era postulated may even have occurred before the link between the Asian
and African francolins was broken since the differences from other francolins shown
by F. gularis, both in ecology and morphology, suggests divergence over a long
period.
CONCLUSIONS
THE GROUPS
The chief lesson to be learnt from comparison of the relationships between the
members of each group re-emphasises the difficulty of assessing from morphological
characters the level of speciation which any form has attained. This is brought
home most forcibly by finding that such dissimilar birds as cranchii and afer in the
Bare-throated Group will interbreed freely, while the three Redwings in the Trans-
vaal shelleyi, levaillantoides, and levaillantii, which look more alike, are able, on
account of ecological divergence, to live in a small area without interbreeding.
It seems that a relatively slight ecological divergence among the francolins may
be effective against interbreeding because their sedentary habits and specialised
choice of habitat make them reluctant to move out of their selected type of vegeta-
tion. These same habits allow also for comparatively small barriers to > act as
isolating factors between two populations that are not necessarily ecologically
divergent. Two instances have been given where rivers appear to form such a
barrier, the Niger between ahantensis and squamatus of the Scaly Group and the
lower Zambezi between natalensis and hildebrandti of the Vermiculated Group.
In the Red-winged Group fifty miles of unsuitable country in Tanganyika between
F. s. shelleyi at Salanga and F. s. uluensis at Mbula serves as barrier to interbreeding
between them.
The difficulty of designating in these circumstances which members of groups
should be called species and which subspecies was discussed at the start of the
paper, for the status of most lies somewhere between the two. This should be
borne in mind in the subsequent discussion on evolution, for these " species ", which
are allopatric members of superspecies, for the most part appear to have attained
a level of speciation far below that of those species which are able to live with each
other in full sympatry.
EVOLUTION
The stages of evolution postulated for each group have been based on what seem
to me to be the simplest and most likely interpretations of the patterns shown, and
they represent the minimum number of stages which can be envisaged to produce
THE FRANCOLINS, A STUDY IN SPECIATION
169
such a pattern. They represent a sequence rather than a time-table of change and
as such could, in theory, be spread over any number of years. However, there is
evidence from the distribution of many montane plants and trees, as well as birds,
which shows that there have been periods in which montane forest has spread
over vast areas of Africa, linking up the present isolated patches in the Cameroons,
Angola, East Africa, Abyssinia, Nyasaland and South Africa, descending as low as
2,000 ft. (see Map 3 A). At the same time (by inference a wet era) there would be
extensive lowland forest below that level. This would have the effect of reducing
the refuges for grassland and woodland species to comparatively small areas isolated
from each other by forests. Such conditions would not only be catastrophic for
many species but would obliterate any traces of the pattern of distribution and
evolution prior to that period. Thus, although such a spread of forest may have
occurred during any extensive cold wet era, it follows that, if the stages of evolution
I have suggested are accepted, the conclusion cannot be evaded that they have
been accomplished since the last spread. This conclusion carries with it implications
on the rate of speciation which need to be examined closely.
In the course of these stages I have suggested that twelve species of francolins
(the ancestral stock of the eight groups and the four independent species) have
increased to forty-one, all the new species being members of superspecies. The
accompanying dendrogram (Fig. 2) summarises the relationship postulated of the
squamatus
ahantensis
griseostriatus
adspersus
capensis
natalensis
hartlaubi
hildebrandti
icterorhynchus
clappertoni
bicalcaratus
bar wood i
swainsonii
rufopictus
leucoscepus
francolinus
pictus
pintadeanus
gularis-
africanus
shelley i
.psilolaemus
levaillantoides
levaillanti
finschi
streptophorus
sephaena
.pondicerianus
Francolinus
FIG. 2. Dendrogram showing the relationship postulated of the various groups and species,
and their date of origin in respect to the last major spread of montane forest.
ZOOL. 10, 2
12
iyo B. P. HALL
various groups and species to each other and their date of origin in respect of this
last spread. Unfortunately no date can be given yet to this last spread of forest,
and there is indeed still uncertainty on the date of onset of the Last Glaciation,
opinions varying between 100,000 and 50,000 years ago. Authorities, however,
agree that it ended about 18,000 years ago and that there have been at least four
glaciations in the course of the Pleistocene. I am suggesting therefore that these
changes have taken place in the comparatively short period of the Late Pleistocene
(i.e. about the last 100,000 years).
If we accept that Francolinus may (among other modern genera of birds) have
originated in the Oligocene (approximately forty to thirty million years ago) (Van
Tyne & Berger 1959 : 6-17) and has taken all the time prior to the last glaciation
to divide into twelve species, it seems that I am postulating an incredible acceleration
in the rate of increase of species. But this is not so, for, as stated, the concept of
periodic spreads of forest over vast areas of Africa is inseparable from the acceptance
of a high casualty rate among the non-forest species at that period. If, for instance,
the present was followed by a cold, wet era that brought with it a wide spread of
forest, it is highly unlikely that any of the less successful non-forest species of
tropical Africa would survive (e.g. rufopictus, streptophorus , finschii, albogularis,
schlegelii) .
Competition between members of different groups in the limited refuges left
would eliminate several of the apparently successful species as well (just as F.
dappertoni appears to have eliminated F. coqui from the Sudan) , while members of
the same group, forced into unprecedented proximity, would probably interbreed
freely. Thus the total number of African species could again be reduced to few
(if any) more than the nine which survived the previous forest spread. These would
be the ancestors of future groups.
Thus while I believe the number of members in any group may fluctuate widely
with successive glacial and inter-glacial eras it seems that any increase in the number
of groups and independent species would be very slow and not incompatible with
the rate of evolution since the inception of the genus.
It could also be suggested that any further increase is unlikely in this figure,
which represents the number of birds which have reached a sufficiently high level of
speciation to be able to live sympatrically without interbreeding, for it is evident
that only a limited number of such birds can be supported in any given area. In
birds like the francolins, in which two species are rarely found actually on the
same ground, this limit is governed largely by the number of alternative types of
habitat, and the unequal distribution of the groups in tropical Africa gives some
reason to suppose that this limit may have been reached.
Disregarding the Montane Group, with its extremely specialised ecology, it will
be seen that, apart from a relict population of the Striated Group, only three groups,
the Scaly, Vermiculated and Red-tailed, are fully represented in the north-western
section (from West Africa to the Sudan and Uganda) . There is little overlap between
the ranges of these groups and where two do occur together one is thriving at the
expense of the other. Furthermore over a vast area it appears that members of the
most successful group, the Vermiculated, have totally driven out representatives of
THE FRANCOLINS, A STUDY IN SPECIATION 171
other groups, namely the red-tailed F. coqui from Lake Chad to the Nile, and the
striated F. streptophorus between the Cameroons and Uganda. This north-western
section is relatively flat with monotonous vegetation, and no very clear-cut divisions
between the vegetation belts : the east and south on the other hand, from Abyssinia
to South Africa and Angola, is more mountainous in many parts with more varied
vegetation, and here no less than six groups are sympatric or partly sympatric.
In this section of Africa I have suggested certain instances in which the range of one
group has been restricted by competition with another, but it is also evident that
several special may thrive in a small area.
I am not competent to discuss the situation in Asia where the development and
status of the francolins must be dependent on those of the numerous other genera
of gamebirds which I have not studied, but it seems unlikely that there is much
room for an increased number of full species.
In arguing that the members of groups may have developed to the level of members
of superspecies within the Late Pleistocene I am conscious that this is postulating
a more rapid rate of speciation than is usually accepted, though Rand (1951: 567)
has suggested that some of the forest subspecies of West Africa may have evolved
in the last 12,000 years, while Moreau (1930) gives examples of Egyptian subspecies
which have probably evolved in 5,000 years. However, some weight is added to
my argument by the evidence collected by Moreau on the climatic fluctuations
that have taken place in this period (Appendix i), which shows that there has been
sufficient variation in both temperature and humidity to bring about the wide
redistributions of the montane and lowland forest, woodland, savanna and steppe
that would be necessary to the different stages proposed. Furthermore, it shows
that all the vegetational changes postulated in the evolutionary sequences in the
groups (which were founded entirely on the evidence of the birds alone) did, or could,
take place, though it is not always easy to satisfy the sequence.
To support some of the major changes which I have envisaged, he has shown, for
instance, that there have been very dry periods prior to 10,000 years ago in Angola
and Katanga (at least) which, if they prevailed elsewhere, could have been sufficient
to connect the northern and southern acacia belts in the way postulated in the
evolutionary sequences of the Bare-throated, Striated and Red-winged Groups. He
gives corroboratory evidence for the existence of such a dry corridor at some period
in Tanganyika, and Benson and White (in press) suggest this corridor may have been
through the Luangwa and Rukwa valleys, and they list many species of birds and
mammals whose present distribution has led them to this conclusion.
Moreau shows also that there was a period (undated) at which Lake Chad was
600 miles long, probably with forest at the southern end joining the main Congo
forest, so forming a barrier between eastern and western woodland or grassland
species, such as was postulated in the Vermiculated Group to split proto-bicalcaratus
from proto-icterorhynchus. He quotes too a record that suggests there was probably
lowland forest at Oldowai in northern Tanganyika in the early Pleistocene where there
is now acacia steppe. Though this dating can hardly be correlated with the spread of
lowland forest envisaged in the later stages of the evolutionary sequence of the
Scaly Group, it suggests that such a spread has taken place once and could
172 B. P. HALL
therefore be repeated in similar climatic conditions: — perhaps achieved in even one
of the recent minor pluvials in East Africa (less than 35,000 years ago).
Similarly he says that late in the Middle Pleistocene there is believed to have been
a dry period when the Congo forest was reduced to strips in the east and west (and
possibly also on the northern rim) and that there was acacia in the country between.
This makes the sequence of evolution postulated for the Red-tailed Group less
improbable than it seemed when I first proposed it (before I had seen Moreau's
paper), though this sequence remains the most difficult to correlate with known
climatic sequences.
Finally, the undated dry period in the Late Pleistocene when the Sahara advanced
300 miles southward may perhaps be correlated with the stage postulated in the
Striated Group when dry conditions prevailed on the ridge of higher country between
the Cameroons and Uganda on which proto-streptophorus may eventually have
been isolated.
All the other climatic changes I have postulated in Africa are relatively small
and the occurrence of similar changes is unquestionable, though not necessarily in
correct sequence. However, some of the dating for these in West Africa is interesting
for the most recent humid period in the north-west (5,000-2,000 years ago) may be
the time at which F. Ucalcaratus spread into Morocco up the north-west coast, and
if so the Morocco population, which shows only slight divergence, may have been
isolated for only 2,000 years. The lack of divergence between the populations of
F. ahantensis on either side of the present gap in the Upper Guinea forest also gives
support to Moreau's evidence that the gap was once further east " probably as
late as the Late Pleistocene ".
In the Spotted Group of Asia, as in the African groups, it was postulated that the
members developed to specific status after the last glaciation. While it is easier
here to argue that the same sequence of isolating and rejoining of species was likely
to take place in any glaciation it seems unlikely that the simple pattern which
remains now could be the product of more than one such sequence.
Finally, lest any of the foregoing should sound dogmatic, it is necessary to end on
the same note of caution that was sounded in the introduction. I am fully aware
that all the conclusions are based on circumstantial evidence drawn from my
interpretations of the picture as we see it today, which are necessarily fallible since
so much of the picture is missing. That it has proved possible to construe the known
facts into a logical and coherent story gives some verisimilitude to it and so does
the evidence that many of the postulated vegetational changes have actually taken
place. However, the story remains hypothetical, but further studies of this nature
and researches in other branches of science will corroborate or disprove others of
the possibilities suggested so that eventually a picture may emerge that approximates
to reality.
SUMMARY
i. It is found that the genus Francolinus (including Pternistis) is comprised of
eight groups, the members of which are largely allopatric, and four independent
species.
THE FRANCOLINS, A STUDY IN SPECIATION 173
2. Some of the difficulties are discussed of denning the relationship between the
members of the groups.
3. The value of different characters as a guide to relationship is illustrated by
comparison between the groups.
4. Each group is studied in turn and the characters and ranges of its component
members defined. In doing this it was found necessary to review the systematics
of the Redwing and Grey wing Francolins of the species africanus (olim. afer),
levaillantoides (olim. gariepensis] and shelleyi.
5. An attempt is made to postulate the sequence of evolutionary stages through
which each group has developed, and the climatic and vegetational changes which
have accompanied each stage.
6. The four independent species are discussed.
7. Comparison between groups suggests that slight ecological divergence may be
an especially effective barrier to interbreeding.
8. It is concluded that most members of groups reflect in their characters,
relationship, and present ranges, the climatic changes that took place in the Late
Pleistocene, subsequent to the last glaciation, or, in Africa, the last extensive spread
of montane forest. Consequently it is suggested that the number of species of this
status is liable to fluctuate considerably in glacial and inter-glacial periods, though
the number of groups and independent species may remain more or less constant.
9. These conclusions are supported to some extent by evidence provided by
R. E. Moreau of climatic changes that have taken place in the Late Pleistocene
(Appendix i), but more evidence is needed from all branches of science to evaluate
them fairly.
APPENDIX i
Summary of some conclusions reached by R. E. Moreau on the climatic fluctuations
in Africa as set out in his paper " Vicissitudes of the African Biota in the Late
Pleistocene " (in press : to be published in Proceedings Zoological Society, London).
Africa generally (in conformity with world temperatures)
Colder than the present (with associated reduced humidity) in varying degrees for
at least 50,000 years prior to 18,000 years ago. In the last extreme phase, extending
from about 24,000-18,000 years ago the cooling amounted to about 5°^- Around
7,000-5,000 years ago probably 2°C. warmer, and for some centuries prior to 150
years ago 1-2° C. cooler, than the present.
Mediterranean Africa and Northern Sahara
A major pluvial period affected at least the western half of the area in the late
Middle Pleistocene, prior to perhaps 100,000 years ago. Palaearctic plants and land-
Footnote : Mr. Moreau has very kindly allowed me to include this summary to illustrate my paper
but it must be appreciated that most of the conclusions quoted have been qualified in the full text, where
reference is given for the authority on which they are based. This summary must not therefore be used
as a base for further premises without consulting the full paper. It must be emphasized that the time-
correlations of the events referred to in different parts of Africa remain to be worked out.
1 74 B. P. HALL
molluscs advanced some 400 miles southward and persisted in Ahaggar until the
Neolithic, some 6,000 years ago.
Between 5,000 and 2,000 years ago it was more humid (at least in the north-west)
than at present.
Southern edge of Sahara and sub-Saharan West Africa
The southern edge of the desert has fluctuated in the course of the Late Pleistocene
(i.e. about the last 100,000 years) between about 300 miles south and 300 miles north
of the present position and these changes probably extended across to the White
Nile. Since the Neolithic, i.e. in about the last 5,000 years, a belt of country some
300 miles in width has reverted to desert.
These fluctuations affected the vegetation of coastal West Africa, especially the
Upper Guinea forest, and when the desert came south of its present location the area
of forest was greatly reduced and further dissected.
At some stage in the Late Pleistocene, not yet dated, Lake Chad increased to an
area 600 miles long, extending nearly to Tibesti. This implies a contemporary north-
ward extension (or movement) of the equatorial rain-belt by some 300 miles.
Prior to its capture by the Lower Niger, some 10,000 years ago, the Upper Niger
expended itself in large lakes northwest of Timbuktu, in what is now extreme desert.
Congo basin and Angola
Late in mid-Pleistocene very dry, with aeolian re-distribution of Kalahari sand.
Congo forest reduced to a strip towards the coast, another along the eastern rim of
the basin and perhaps a third along the northern edge. Throughout Late Pleistocene,
humid except in Katanga and Eastern Angola. Here arid in the latter part of the
Late Pleistocene down to about 10,000 years ago.
East Africa
The " Gamblian " pluvial, with fluctuations in intensity was contemporary with
the Last Glaciation and extended down to about 18,000 years ago, and there have
been subsequently at least two short periods more humid than the present. Owing
to the broken topography the vegetational effects would have been, in the main,
local advances and retractions of vegetation types. In Somaliland the fluctuations
were minor and the country always to some degree dry. (Note : There is evidence
that at some period in the Early Pleistocene lowland forest extended at least to
Oldowai in northern Tanganyika. Also at some period in the Late Pleistocene a
corridor of acacia probably extended through Tanganyika to Northern Rhodesia).
Rhodesia and southern Africa
During the Late Pleistocene the rainfall fluctuated between about half and one and
a half times the present in the Rhodesias and the Transvaal. Kalahari sand was
blowing east as far as the Victoria Falls some 12,000-9,000 years ago. The Orange
Free State was drier than the present some 40,000 years ago. South West Africa was
never better than semi-arid,
THE FRANCOLINS, A STUDY IN SPECIATION 175
APPENDIX 2
Taxonomic notes and synonymy
SPOTTED GROUP
F. francolinus
Five clinal races have been recognised and further sub-division does not seem useful. There-
fore caucasicus and sarudyni are placed in the synonymy of francolimts ; festinus Koelz, 1955
and bogdanovi in the synonymy of henrici ; parkerae Van Tyne & Koelz, 1955 in the synonymy
of asiae. Also billypaynei Meinertzhagen, 1933, which is a name given to an exceptionally dark
micro-population from the Lake of Antioch, is included in francolinus.
F. pintadeanus
The characters on which wellsi was based do not seem constant and it is accordingly regarded
as a synonym of pintadeanus.
I accept Delacour's opinion (1951 : 10) that -F. boineti Bourret, 1944 is an aberrant specimen of
F. pintadeanus.
BARE-THROATED GROUP
F. leucoscepus
The amount of variation exhibited by the whole species is so slight that numerous sub-
divisions are not practical. Accordingly holtemulleri, muhamedbenabdullah, keniensis,* kili-
mensis, tokora, oldowai Van Someren, 1939, are placed in the synonymy of infuscatus.
F. swainsonii
Since Pternistis has been submerged in Francolinus, P. s. chobiensis Roberts, 1932, is pre-
occupied and F. s. lundazi used instead (see Benson & White, 1957: 138).
A record of the occurrence of F. swainsonii at Karonga, at the north end of Lake Nyasa
(Long, 1961 : 25) was based on a misidentification, the specimens being F. afer (Benson,
in lift.).
F. afer
In the block of vermiculated subspecies there is much individual but little geographical
variation. Accordingly punctulata and nyanzae are placed in the synonymy of cranchii.
In the black-and-white birds of Angola and South West Africa variation between the northern
and southern birds is too slight to warrant recognition, so palliditectus White, 1958 (nom. nov.
for cunenensis Roberts) is placed in the synonymy of afer. In South Africa an intermediate
population between castaneiv enter and notatus was named krebsi, which I place in the synonymy
of castaneiv enter. In eastern Africa the name humboldtii was given to a bird from Tete on the
Zambezi, a locality lying in the centre of the zone of hybrids between the black-faced melano-
gaster and the white-faced swynnertoni : it is here restricted to the unstable population.
If names are required for the hybrids between the vermiculated and black-and-white sub-
species it is convenient in Angola to refer those populations which are closest to cranchii to
manueli White, 1945 (synonyms camabatelae Meise, 1958, and mackenziei White, 1945). an^
those which are closest to afer to benguellensis (synonyms angolensis Bocage, angolensis Roberts,
and chio Meise, 1958 — see Traylor ig6oa: 143-146). Similarly in Tanganyika, Nyasaland and
the Luangwa valley those populations closest to intercedens may be known as bohmi (synonym
tornowi Meise, 1933), and those closest to melanogaster may be known as itigi (synonym
aylwinae White, 1947).
With the merging of Pternistis in Francolinus F. coqui angolensis Rothschild, 1902, is ante-
dated by Pternistes afer var. angolensis Bocage, 1893, and by the laws of homonymy should be
renamed. However Bocage 's name angolensis was given in error to the same bird which he had
named benguellensis at the head of his article, and the name was forgotten until listed in the
synonymy of Pternistis afer benguellensis by Bowen (1930: 15?) • Jt thus has remained unused
as a senior synonym in the primary zoological literature for more than fifty years and must
be considered a forgotten name (nomen oblitum). (Int. Code of Zool. Nomen. 1961. Article
* Also pre-occupied by F. squnmatus keniensis Mearns, 1910.
176 B. P. HALL
23b.) Application is being made to the International Commission for it to be placed on the
Official Index of Rejected Names.
MONTANE GROUP
F. castaneicollis
It is probable that most of the micro-populations in isolated patches of evergreen forest are
slightly distinct, but it is not practical to recognise by name all those that show minor
differences: accordingly vottegi and gofanus are placed in the synonymy of castaneicollis, and
patrizii Toschi, 1958, in the synonymy of kaffanus.
F. jacksoni
It has not been established that variation in the amount of white in the underparts is geographi-
cal or associated with altitude. Therefore gurae is placed in the synonymy of jacksoni.
F. nobilis
F. camerunensis ruandae Van Saceghem, 1942, is placed in the synonymy of F. n. nobihs.
It is also pre-occupied by -F. coqui ruandae Jackson, 1938.
F. swierstrai
F. cvuzi Themido, 1935, is placed in the synonymy of F. swierstrai.
SCALY GROUP
F. squamatus
Although there is considerable individual variation no significant geographical variation is
recognised from the greater part of the range. Accordingly tetraoninus, zappeyi and dowashanus
are placed in the synonymy of schuetti and whitei Schouteden 1954 (nom. nov. for confusa
Schouteden 1954) *n tne synomy of squamatus.
It is not thought practical to recognise by name any micro-populations isolated on the
mountains of the Kenya/Tanganyika border, since the main trend of variation is clinal.
Therefore, kapitensis, keniensis and chyuluensis Van Someren, 1939 are placed in the synonymy
of maranensis.
VERMICULATED GROUP
F. bicalcaratus
It has been shown that birds become darker and more heavily patterned towards the south-
east of the range, but the differences exhibited in this cline do not seem great enough to warrant
separating the birds of the extreme south-east from those of the southern (British) Cameroons.
Accordingly molunduensis Grote, 1949, is synonymised with ogilviegranti.
F. icterorhynchus
The clinal variation does not warrant more than a division into two subspecies. Accordingly
grisescens is placed in the synonymy of icterorhynchus and emini and ugandensis in the synonymy
of dybowskii. Birds showing a few chestnut feathers on the flanks, the character on which
ugandensis was founded, occur sporadically in south-eastern Uganda and must be considered
aberrant specimens perhaps indicating past hybridisation with clappertoni.
F. clappertoni
F. tschadensis is placed in the synonymy of F. c. clappertoni (see text).
The clinal variation shown in southern populations does not warrant extensive splitting;
accordingly cavei Macdonald, 1940, and testis are placed respectively in the synonymy of gedgii
and sharpii.
F. hildebrandti
Numerous subdivisions are not practical; accordingly helleri is placed in the synonymy of
hildebrandti, and grotei and lindi in that of johnstoni.
F. natalensis
The name thamnobium Clancey, 1953, was proposed to distinguish paler birds inhabiting drier
areas from those of wetter habitats in Natal and the Zoutspansberg (natalensis), but since this
THE FRANCOLINS, A STUDY IN SPECIATION 177
gives natalensis a discontinuous range it is not practical to recognise the differences by name.
F. hartlaubi
ovambensis is considered a synonym of bradfieldi.
F. adspersus
kalahari is considered a synonym of adspersus.
STRIATED GROUP
F. sephaena
The name somaliensis Grant & Praed, 1934, was given to streaked birds from Somaliland
and was based on the misapprehension that the type of spilogaster was an unstreaked bird
(Rand, 1950: 384). As noted in the text I believe spilogaster must be used for the hybrid
populations from Somaliland to southern Kenya. However, if it is established that there still
exists an unmixed population of streaked birds on the coastal plains which is distinct from the
southern rovuma then the name somaliensis is available. Such a population can be assumed
to have existed once since it must be regarded as one of the parents of the hybrid spilogaster
but it may have been integrated into the hybrid population.
Local variations in size, colour and pattern throughout the species do not seem great enough
to distinguish by name. Accordingly zuluensis is placed in the synonymy of sephaena :
thompsoni, chobiensis, mababiensis in the synonymy of zambesiae : schoanus, ochrogaster Hartlaub,
1882, delutescens, and jubaeensis in the synonymy of grantii: Peters (1934: 73) shows that
rovuma antedates kirkii.
RED-WINGED GROUP
F. psilolaemus
On the author's own showing fricki is regarded as a synonym of F. p. ellenbecki (Friedmann,
1930: 113).
F. shelleyi
There is not sufficient variation between the populations of South Africa and Tanganyika
to justify the recognition of trothae which is regarded as a synonym of shelleyi, as is sequestris
Clancey, 1960.
F. africanus
Praed & Grant have been among the few modern authors who have correctly regarded
Perdix afra Latham, 1790, as the senior name of the South African Grey wing, known by others
as F. africanus Stephens, 1890. However, since the submersion of Pternistis in Francolinus,
Perdix afra Latham is pre-occupied by Tetrao afer Muller 1776, the senior name for the bare-
throated francolin formerly known as Pternistis afer. The Greywing must therefore now be
known as F. africanus.
Within the range of the species there is slight local variation but it is not great enough for
me to recognise any subspecies: proximus Clancey, 1957, i-s therefore placed in the synonymy.
F. levaillantoides
Roberts (1936: 321) shows that F. levaillantoides is the original name for this species and
gariepensis is a synonym.
In this species the haphazard distribution of rufous and grey micro-populations in the
southern districts of the Sudan and Abyssinia and in Angola and South West Africa makes it
difficult to know how many subspecies can usefully be recognised. I believe it is impractical
to attempt to differentiate between these variations if no other differences are shown, and,
furthermore, names should not be used for intermediate populations. Accordingly I recognise
in the south —
(a) F. I. levaillantoides. All districts east and south of the Kalahari, intergrading with
kalaharica on the fringes of the Kalahari. Richly coloured (even in the greyest specimens)
and heavily patterned. Synonyms, ludwigi and gariepensis.
(b) F. I, kalaharica (Roberts) 1932. The Kalahari, Pale and grey, birds from central
178 B. P. HALL
districts almost unmarked on the abdomens; birds from the fringes rather darker with
more markings on the abdomen. Synonym, langi Roberts, 1932. (Although langi has
line priority over kalaharica I use kalaharica, as I am entitled to do as first revisor, since
the name was given to a very pale, grey bird from the Damara Pan near the centre of the
Kalahari, and thus represents the extreme of variation, while langi was given to a rather
darker bird from Nkate near the eastern border.)
(c) F. I. pallidior. South West Africa. Either rufous or grey but the rufous birds paler in
tone than levaillantoides, and the grey birds darker and more heavily marked than
kalaharica. As known at present the population of the Windhuk area is more rufous,
and that of the Tsumeb area greyer (approaching kalaharica). Synonym wattii
Macdonald, 1953. (See Macdonald, 1953, for detailed discussion.)
(d) F. I. jugularis. Southern Angola to the Cunene. Differs from all other subspecies in
having a broad black-and-white gorget: typical birds from the Benguela area are small
and grey but those from the Cunene are larger and more rufous showing an approach to
some of the populations of pallidior. Synonyms, cunenensis Roberts, 1932, stresemanni
Hoesch & Niethammer, 1940.
In the north —
(e) F. 1. archeri. Extreme southern Sudan to the Golis range of western (British) Somaliland.
Similar to F. I. pallidior in having both rufous and grey populations but in both the
abdomens are paler and commonly marked with fine lines. Synonyms friedmanni
Grant & Praed, 1934, stantoni Cave, 1940.
(/) F. I. lorti. Mountains of eastern (British) Somaliland. Very grey, with a very pale
abdomen fairly heavily marked with fine lines.
(g) F. I. gutturalis. Northern Abyssinia and Eritrea. Quite distinct (see text). Synonym
eritrea Zedlitz.
F. levaillantii
A number of names have been given to the birds of the northern and western populations
which have no black-and-white stripe down the ochre collar on the hind neck. However,
while the few specimens available indicate that there is a tendency for Angola birds to be
paler, the difference is trivial and the series otherwise is remarkably uniform. Accordingly
mulemae, adolffriederici, benguellensis Neumann, 1908, clayi White, 1944, and momboloensis
White, 1952 (nom. nov. for benguellensis) are placed in the synonymy of kikuyuensis .
THE RED-TAILED GROUP
F. coqui
In the populations of central and south-eastern African there is considerable individual and
local variation in size and colour so that consistent geographical variation is hard to determine.
I do not believe it is practical to distinguish from nominate coqui the small coastal birds of
Kenya since equally small birds are found in Natal, or to distinguish the populations of Natal,
Zululand and southern Mozambique, which on the whole are less rufous, for individuals can be
matched with others elsewhere. The names stuhlmanni and campbelli are therefore placed in
the synonymy of coqui. I regard lynesi Sclater, 1932, also as a synonym of coqui since the type
is intermediate between coqui and kasaicus but comes from an area in which the majority of
specimens are closest to coqui.
For the Uganda race the original spelling " ruahdae " has been used since, while the author
may have intended the name to be " ruandae ", there is no clear evidence of this intention in
the original publication. (Int. Code of Zool. Nomen. 1961, Article 32a (ii) .) F. coqui ruandae
Jackson, 1938, is accordingly a junior objective synonym. (Article 33a (ii) .)
F. albogularis
Serle (1957: 388) shows that gambagae must be considered a synonym of buckleyi.
The isolated populations of Marungu and Upemba in the south-eastern Congo, the Balovale
district of Northern Rhodesia, and of eastern Angola seem to show an increase from east to
west in the amount of patterning in the females, and the Angola birds seem also to be darker,
THE FRANCOLINS, STUDY IN SPECIATION 179
less rufous, than others (Hall, ig6oa: 409). Too few specimens are available at present on
which to judge the extent of variation, and until there are more, especially of topotypical
meinertzhageni from Rhodesia, it seems best to use meinertzhageni for Rhodesian and Angola
birds and dewittei for the Congo birds, though I believe that the Rhodesian birds may prove to
be closer to those from the Congo than to those from Angola.
F. schlegelii
Traylor (1960: 86) has shown that confusus Neumann, 1933, should be considered a synonym
of schlegelii.
SPECIES NOT INCLUDED IN ANY GROUP
F. pondicerianus
It has been shown that variation in this species is slight and largely clinal and ecological.
It is not practical to recognise more than one intermediate subspecies. Accordingly paganus
Koelz, 1954, ti*ar Koelz, 1954, prepositus Koelz, 1954, are au placed in the synonymy of
interpositus.
SPOTTED GROUP— MAP 1
The shaded areas represent the overall ranges of the species. The thick red line indicates
divisions between species. The thin red line indicates divisions between subspecies, the
zig-zag line denoting intergradation. The red crosses indicate localities from which
hybrid specimens have been obtained.
F. francolinus
F. pictus
1. francolinus
2. arabistanicus
3. henrici
4. asiae
5. melanotus
6. pallidus
7. pictus
8. watsoni
F. pintadeanus 9. phayrei
10. pintadeanus
MAP. IA. Hypothetical distribution of the ancestral stock of the three species during a
glaciation (Stage 2).
BARE- THROATED GROUP— MAP 2
Overall ranges are shown rather than precise collecting localities. Thick red lines
denote specific boundaries. Thin red lines denote subspecific boundaries, plain lines
being boundaries between non-intergrading subspecies, zig-zag lines boundaries between
intergrading subspecies, toothed lines boundaries of hybrid zones. The red " X " marks
the area of hybrids swainsonii x afer. Numbers have been given only to species and
subspecies which are important to the discussion on speciation, but the ranges of other
subspecies are indicated.
i. F. leucoscepus (leucoscepus, infuscatus)
iiillii-iiililiiliil 2- F. rufopictus
V.V.'.'.'.'.J 3. F. swainsonii (swainsonii, lundazi, damarensis, gilli)
IB
F. afer. Black-and-white, " afer-type " subspecies
4. castaneiv enter & notatus
5. lehmanni
6. swynnertoni
6x7. hybrids (humboldtii)
7. melanogaster & loangwae
8. leucoparaeus
9. afer
Vermiculated, " cranchii-type " subspecies
10. cranchii & intercedens
11. harterti
Hybrids between " afer-type " and " cranchii-type " subspecies.
MAP 2A. Hypothetical distribution of the ancestral stock of the four species during a
spread of forest (Stage 4).
MAP 2B. Hypothetical distribution of the ancestral stock of the four species and two main
blocks of afer subspecies during a later dry period (Stage 6) .
MONTANE GROUP— MAP 3
The letters indicate precise localities from which specimens have been collected.
Plain red lines denote divisions between isolates which are regarded as species, and
broken lines between those regarded as subspecies. The thickness of the lines is relative
to the degree of divergence between neighbouring forms.
e F. erckelii i. pentoni
2. erckelii
o F. ochropectus 3. ochropectus
c F. castaneicollis 4. ogoensis
5 . castaneicollis
6. kaffanus
7. atrifrons
j F. jacksoni 8. jacksoni
g. pollenorum
n F. nobilis 10. nobilis
1 1 . chapini
x F. camerunensis 12. camerunensis
s F. swierstrai 13. swierstrai
MAP 3A. Hypothetical distribution of montane forest (shaded black) in a cold, wet era
(Stage i).
ZOOL. 10, 2.
SCALY GROUP— MAP 4
A continuous distribution has been shown for members of the Scaly Group through the
Upper and Lower Guinea forests, along the escarpment of Angola, and in the wetter areas
of Uganda and Kenya, for within these areas the birds may be expected in suitable
clearings or cultivations. Outside this range only general localities from which specimens
have been collected have been shaded. Broad red lines indicate divisions between species :
thinner red lines denote divisions between subspecies, those that intergrade being indicated
by a zig-zag line. Thicknesses of the subspecific lines are relative to the divergence shown
by neighbouring forms.
F. ahantensis
1. hopkinsoni
2. ahantensis
F. squamatus
- griseostriatus
3. squamatus
4. schuetti
5. maranensis
6. usambarae
7. uzungwensis
8. doni
MAP 4A. Hypothetical map showing extent of lowland forest in a warm, wet era (Stage 5).
Limits of forest shown by broken line.
ZOOL. IO, 2
VERMICULATED GROUP— MAP 5
The distribution of many members of this group is known to be discontinuous.
Accordingly only general areas from which specimens have been collected are shaded,
but the birds may be expected to occur in some of the intervening areas in which little
collecting has been done. Thick red lines denote divisions between species: where there
is evidence of hybridisation these lines are toothed. Thin red lines denote divisions
between subspecies, those that intergrade being shown with a zig-zag line. The red
" X " denotes a hybrid between hildebrandti and natalensis. The red " T " denotes the
aberrant or hybrid specimen F. tschadensis. The red " U " denotes the area in which
the aberrant " ugandensis "-type birds are found.
F. bicalcaratus i. ayesha
2. bicalcaratus
3. adamauae
4. ogilviegranti
5. thornei
F. clappertoni 6. clappertoni
8. heuglini
9. sharpii
10. konigseggi
11. nigrosquamatus
F. icterorhynchus 12. dybowskii
13. icterorhynchus
F. hildebrandti
F. natalensis
F. hartlaubi
F. capensis
F. adspersus
F. harwoodi
14. altumi
15. hildebrandti
1 6. johnstoni
17. neavei
1 8. natalensis
(hartlaubi, bradfieldi, crypticus}
tfll
STRIATED GROUP— MAP 6
Only general areas from which birds have been collected are shaded. The broken thick
red line indicates the outer limits of the range of F. streptophorus , and the unbroken thick
red line the outer limits of the range of F. sephaena. Thin red lines indicate subspecific
divisions, plain lines denoting divisions without intergrading, zig-zag lines denoting
intergrading, and toothed lines denoting the boundary of a hybrid area.
F. streptophorus
F. sephaena i. sephaena
2. zambesiae
3. grantii
3x4. spilogaster
4. rovuma
MAP 6A. Hypothetical distribution of the ancestral stock of the two species at the end
of a dry era (Stage 4). Arrows indicate the line of advance of encroaching woodlands.
«,vif
RED -WINGED GROUP— MAP 7
The distribution of all members of this group is discontinuous, accordingly only general
areas from which birds have been collected are shaded. Thick red lines denote the
extreme limits of the range of each species : these lines are broken where they link remotely
isolated populations, except in F. levaillantoides where arrows indicate where the lines
linking the northern and southern populations might fall. Lineal divisions between
subspecies are not shown but different shading has been used for those subspecies which
are strongly differentiated: others are numbered. The red cross indicates the Amani
specimen which is intermediate between F. s. shelleyi and F, s. uluensis.
F. psilolaemus (psilolaemus, ellenbecki, elgonensis, theresae]
F. shelleyi uluensis (including macarthuri)
F. shelleyi shelleyi
F. shelleyi whytei
F. africanus
F. levaillantoides i. levaillantoides
2. kalaharica
3. pallidior
4. jugularis
5. archeri
6. lorti
F. levaillantoides gutturalis
F. levaillantii
F. finschi
7. levaillantii
8. crawshayi
g. kikuyuensis
MAP yA. Mean Annual Rainfall (taken from Fitzgerald 1950: 35)
under 4 inches
MAP ye. Diagrammatic map showing hypothetical colonisation routes and centres of
speciation (Stages 3-6).
ad
RED-TAILED GROUP— MAP 8
The distribution of all members of this group is discontinuous and only general areas
from which specimens have been collected are shaded. Thick red lines denote the extreme
limits of the ranges of species, those linking remotely isolated populations being broken,
except between the north-western and southern populations of F. albogularis where the
link is indicated by arrows. Subspecific divisions are shown lineally only in the critical
area of Kenya and Tanganyika. Here a toothed line surrounds a population which
appears to be a result of hybridisation but which does not now intergrade with neigh-
bouring subspecies: zig-zag lines indicate intergrading. Outside this critical area all
subspecies, if not isolates, intergrade.
F. coqui
W///A
F. schlegelii
F. albogularis
1. coqui
2. vernayi
3. hoeschianus
4. angolensis
5. kasaicus
6. ruahdae
7. hubbardi
8. thikae
9. maharao
10. spinetorum
11. albogularis
12. buckleyi
13. dewittei
14. meinertzhageni
MAP SA. Hypothetical distribution of the ancestral stock of F. coqui and proto-albogularis/
schlegelii (shown by mixed diagonal and horizontal shading) before the two latter species
diverged (Stage 3). Distribution of lowland forest shown in black.
MAP SB. Hypothetical distribution of the ancestral stock of the three species in a sub-
sequent dry era (Stage 4).
SPECIES NOT INCLUDED IN ANY GROUP
MAP 9
F. nahani
F. lathami
1. lathami
2. schubotzi
D
F. pondicerianus
F. gularis
MAP 10
1. mecranensis
2. interpositus
3. pondicerianus
4. ceylonensis
10
VEGETATION MAP OF AFRICA— MAP 11
Adapted from " Vegetation Map of Africa south of the Tropic of Cancer", Oxford
University Press, 1959.
The numbering and terminology of the original map have been used as far as possible,
excluding Madagascar. Explanatory notes indicating geographical areas are given in
the key in cases where it is not possible to differentiate on the map between vegetation
types.
KEY
1-4. EVERGREEN FORESTS & MONTANE COMMUNITIES.
1. Montane Evergreen Forest.
2. Temperate and Subtropical Evergreen Forest. Knysna and other
districts of the extreme south only,
3. Montane Communities — undifferentiated. These include evergreen
forest (type i), grassland (type 5) and woodland.
4. Montane Communities — with alfro-alpine communities. High moun-
tains of central and east Africa and Ethiopia only.
(It has not been possible on this scale to distinguish between types i ,
3, and 4.)
5-6. MONTANE & HIGH ALTITUDE GRASSLAND.
5. Montane Grassland above 6,000 ft. North-eastern and central Africa.
6. Temperate and Subtropical Grassland. Pure grassland above 3,5oo//.
in South Africa.
MOIST FOREST AT LOW AND MEDIUM ALTITUDES.
FOREST-SAVANNA MOSAIC.
8. Moist forest surrounded by savanna of tall grasses. Found in belts
surrounding type 7.
9. Coastal Forest-Savanna Mosaic. East coast only.
10. DRY & DECIDUOUS FOREST — with abundant Baikiaea plurijuga. Found
only in parts of N. & S. Rhodesia, Bechuanaland and the Caprivi Strip,
and sometimes regarded as woodland rather than forest.
12 & 14. THICKETS.
12. Itigi type. Central Tanganyika only.
14. Ethopian evergreen type. Ethiopia only.
15. CAPE MACCHIA.
16. RELATIVELY MOIST WOODLANDS. Types in which the genera
Isoberlinia, Brachystegia, and Julbernadia are absent or rare.
17-19. WOODLANDS (interspersed with Savanna).
17. With abundant Isoberlinia doka and /. dalzielii. Northern areas.
18. With abundant Brachystegia and Julbernadia — " myombo ". Tan-
ganyika to central Northern Rhodesia, Southern Rhodesia and Mozambique.
19. Similar to type 18 but with a number of distinctive species. Prin-
cipally on Kalahari sand. Angola to western N. Rhodesia.
20-22. RELATIVELY DRY WOODLANDS & SAVANNAS.
20. With frequent Acacia but other species as well, and savannas of tall
grass with certain species of Acacia.
21. Ethiopian types. A heterogeneous assemblage of low deciduous
shrubs on stony ground. Ethiopia only.
22. With abundant Colospermum mopane. Southern Africa only.
24. GRASS STEPPE ON KALAHARI SAND.
25. STEPPES.
25. Wooded steppe with abundant Acacia and Commiphora.
26. Grass steppe with thicket clumps. Western Uganda only.
27. Grass steppe — Luanda type. Coastal Angola only.
28-31. KAROO & SUBDESERT STEPPE.
28. Karoo Succulent Steppe. Western Cape Province only.
29. Karoo shrub and grass. Cape Province and southern South West Africa.
30. Transitional and mixed Karoo. East-central Cape Province.
31. Northern areas and northern South West Africa and southern Angola.
32. DESERT.
33- SWAMPS.
200 B. P. HALL
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THE FRANCOLINS, A STUDY IN SPECIATION 201
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INDEX TO SPECIFIC AND SUBSPECIFIC NAMES OF FRANCOLINUS
Specific and subspecific names of francolins are listed under the nomenclature used in this
paper, those of species and subspecies here recognized appearing in roman type, others in italics.
No attention is paid to other generic names that have been given to francolins except in a
few cases where the merging of these names in Francolinus has created homonyms.
For any species that is here regarded as a member of a species group all the pages of the
section dealing with the whole group are listed, regardless of whether the species is actually
mentioned on each page.
adamauae, F. bicalcaratus, 144, 188 afra, Perdix (F. afer), 152, 155, 173, 177
adolffriederici, F. levaillantii, 178 africanus, F., no, in, 117, 120, 152-160,
adspersus, F., no, 113, 117, 120, 121, 140- 169, 173, 177, 192
148, 169, 176, 188 ahantensis, F., no, in, 116, 136-140, 167,
afer, F., no, in, 113, 116, 119, 121, 122, 168, 169, 172, 186
125-131, 132, 140, 145, 146, 168, 169, 175, ahantensis, F. a., [136], 186
177, 182 albogularis, F., no, 114, 117, 122, 160-165,
afer, F. a., 127, 175, 182 169, 170, 178, 194
202
B. P. HALL
albogularis, F. a., 162, 194
altumi, F. hildebrandti, 142, 146, 188
angolensis, Pternistis afer, 175
angolensis, F. coqui, 163, 175, 194
arabistanicus, F. francolinus, 124, 180
archeri, F. levaillantoid.es, 155, 156, 178, 192
asiae, F. francolinus, 124, 175, 180
atrifrons, F. castaneicollis, 112, 133, 135, 136,
184
ayesha, F. bicalcaratus, 141, 144, 147, 188
aylwinae, F. afer, 175
benguellensis , F. after, 175
benguellensis, F. levaillantii, 178
bicalcaratus, F., no, 117, 118, 120, 121, 123,
140-148, 161, 164, 165, 169, 171, 172, 176,
1 88
bicalcaratus, F. b., 144, 188
billypaynei, F. francolinus, 175
bogdanovi, F. francolinus, 175
bohmi, F. afer, 175
boineti, F., 175
bottegi, F. castaneicollis, 176
bradfieldi, F. hartlaubi, 145, 177, 188
buckleyi, F. albogularis, 162, 178, 194
camabatelae, F. afer, 175
camerunensis, F., no, 116, 118, 120, 131-136,
169, 184
campbelli, F. coqui, 178
capensis, F., no, 117, 140-148, 169, 188
castaneicollis, F., no, in, 116, 131-136, 169,
176, 184
castaneicollis, F. c., 133, 135, 176, 184
castaneiventer, F. afer, 128, 130, 131, 175, 182
caucasicus, F. francolinus, 1 75
cavei, F. clappertoni, 176
ceylonensis, F. pondicerianus, 167, 196
chapini, F. nobilis, 134, 184
chio, F. afer, 175
chobiensis, F. sephaena, 177
chobiensis, Pternistis swainsonii, 175
chyuluensis, F. squamatus, 176
clappertoni, F., no, 114, 117, 118, 119, 120,
121, 140-148, 161, 164, 165, 169, 170, 176,
188
clappertoni, F. c., 141, 144, 176, 188
clayi, F. levaillantii, 178
confusa, F. squamatus, 176
confusus, F. schlegelii, 179
cooperi, Pternistis, 128
coqui, F., no, in, 117, 118, 119, 121, 122,
145, 147, 160-165, 167, 169, 170, 171, 178,
194
coqui, F. c., 161, 162, 163, 164, 178, 194
cranchii, F. afer, 127, 168, 175, 182
crawshayi, F. levaillantii, 157, 192
cruzi, F., 176
crypticus, F. hartlaubi, 145, 188
cunenensis, F. levaillantoides, 178
cunen sis, Pternistis afer, 175
damarensis, F. swainsonii, 129, 182
delutescens, F. sephaena, 177
dewittei, F. albogularis, 162, 179, 194
doni, F. squamatus, 137, 139, 186
dowashanus, F. squamatus, 176
dybowskii, F. icterorhynchus, 144, 176, 188
elgonensis, F. psilolaemus, 153, 154, 192
ellenbecki, F. psilolaemus, 153, 177, 192
emini, F. icterorhynchus, 176
erckelii, F., no, in, 116, 120, 131-136, 141,
143, 145, 146, 169, 184
erckelii, F. e., 135, 184
eritrea, F. levaillantoides, 178
festinus, F . francolinus , 175
finschi, F., no, 117, 119, 122, 152-160, 169,
170, 192
francolinus, F., no, in, 116, 118, 119, 122,
123-125, 165, 166, 167, 168, 169, 175, 180
francolinus, F. 1, 124, 175, 180
fricki, F. psilolaemus, 177
friedmanni, F. levaillantoides, 155, 178
gambagae, F. albogularis, 178
gariepensis, F., 155, 173, 177
gedgii, F. clappertoni, 144, 176, 188
gilli, F. swainsonii, 129, 182
gofanus, F. castaneicollis, 176
grantii, F. sephaena, 150, 151, 152, 177, 190
grisescens, F. icterorhynchus, 176
griseostriatus, F., no, 116, 136-140, 167, 169
grotei, F. hildebrandti, 176
gularis, F., 109, no, 117, 167-168, 169, 196
gurae, F.jacksoni, 176
gutturalis, F. levaillantoides, 155, 156, 159,
160, 178, 192
harterti, F. afer, 127, 182
hartlaubi, F., no, 117, 120, 121, 140-148,
169, 177, 188
hartlaubi, F. h., 144, 188
harwoodi, F., no, 117, 118, 120, 140-148,
169, 1 88
helleri, F. hildebrandti, 176
henrici, F. francolinus, 124, 175, 180
THE FRANCOLINS, A STUDY IN SPECIATION
heuglini, F. clappertoni, 144, 188
hildebrandti, F., no, in, 117, 118, 140-148,
1 68, 169, 176, 1 88
hildebrandti, F. h., 142, 176, 188
hoeschianus, F. coqui, 163, 194
holtemulleri, F. leucoscepus, 175
hopkinsoni, F. ahantensis, 136, 186
hubbardi, F. coqui, 161, 162, 164, 194
humboldtii, F. afer, 127, 175, 182
icterorhynchus, F., no, 114, 117, 120, 122,
140-148, 163, 164, 165, 169, 171, 176, 188
icterorhynchus, F. i., 144, 176, 188
infuscatus, F. leucoscepus, 129, 175, 182
intercedens, F. afer, 127, 175, 182
intermedius, F., 124
interpositus, F. pondicerianus, 167, 179, 196
itigi, F. afer, 175
jacksoni, F., no, 116, 118, 120, 131-136, 169,
176, 184
jacksoni, F. j., 133, 176, 184
johnstoni, F. hildebrandti, 142, 176, 188
jubaeensis, F. sephaena, 177
jugularis, F. levaillantoides, 156, 178, 192
kaffanus, F. castaneicollis, 133, 134, 135, 176,
184
kalahari, F. adspersus, 176
kalaharica, F. levaillantoides, 156, 177, 178,
192
kapitensis, F. squamatus, 176
kasaicus, F. coqui, 163, 178, 194
keniensis, F. leucoscepus, 175
keniensis, F. squamatus, 175, 176
kikuyuensis, F. levaillantii, 157, 178, 192
kilimensis, F. leucoscepus, 175
kirkii, F. sephaena, 177
konigseggi, F. clappertoni, 144, 188
krebsi, F. afer, 175
langi, F. levaillantoides, 178
lathami, F., 109, no, 117, 119, 123, 165-166,
169, 196
lathami, F. 1., [166], 196
lehmanni, F. afer, 128, 131, 182
leucoparaeus, F. afer, 127, 131, 182
leucoscepus, F., no, 113, 116, 119, 122, 125-
131, 169, 175, 182
leucoscepus, F. 1., 129, 182
levaillantii, F., no, 117, 121, 122, 152-160,
168, 169, 178, 192
levaillantii, F. 1., 157, 192
203
levaillantoides, F., no, 113, 117, 121, 122,
152-160, 168, 169, 173, 177, 192
levaillantoides, F. 1., 156, 177, 178, 192
lindi, F. hildebrandti, 176
loangwae, F. afer, 127, 129, 182
lorti, F. levaillantoides, 155, 156, 178, 192
ludwigi, F. levaillantoides, 177
lundazi, F. swainsonii, 129, 175, 182
lynesi, F. coqui, 178
mababiensis , F. sephaena, 177
macarthuri, F. shelleyi, 154, 192
mackenziei, F. afer, 175
maharao, F. coqui, 161, 164, 165, 167, 194
manueli, F. afer, 715
maranensis, F. squamatus, 137, 176, 186
mecranensis, F. pondicerianus, 167, 196
meinertzhageni, F. albogularis, 162, 179, 194
melanogaster, F. afer, 127, 129, 131, 175, 182
melanotus, F. francolinus, 124, 180
molunduensis, F. bicalcaratus, 176
momboloensis, F. levaillantii, 178
muhamedbenabdullah, F. leucoscepus, 175
mulemae, F. levaillantii, 178
nahani, F., 109, no, 117, 120, 123, 166-167,
169, 196
natalensis, F., no, 111, 117, 121, 128, 131,
140-148, 168, 169, 176, 188
natalensis, F. n., 117, 142, 176, 177, 188
neavei, F. natalensis, 117, 142, 143, 146, 188
nigrosquamatus, F. clappertoni, 144, 188
nobilis, F., no, 116, 120, 131-136, 169, 176,
184
nobilis, F. n., [134], 176, 184
notatus, F. afer, 128, 131, 175, 182
nyanzae, F. afer, 175
ochrogaster, F. sephaena, 177
ochropectus, F., 107, no, in, 112, 116, 131-
136, 169, 184
ogilviegranti, F. bicalcaratus, 144, 176, 188
ogoensis, F. castaneicollis, 133, 184
oldowai, F. leucoscepus, 175
ovambensis, F. hartlaubi, 177
paganus, F. pondicerianus, 179
pallidior, F. levaillantoides, 156, 178, 192
palliditectus, F. afer, 175
pallidus, F. pictus, 124, 180
parkerae, F. francolinus, 175
patrizii, F. castaneicollis, 176
pentoni, F. erckelii, 132, 135, 145, 184
204
B. P. HALL
phayrei, F. pintadeanus, 124, 180
pictus, F., no, in, 116, 119, 120, 123-125,
167, 169, 1 80
pictus, F. p., 124, 1 80
pintadeanus, F., no, 116, 118, 123-125, 169,
175, 180
pintadeanus, F. p., [124], 175, 180
pollenorum, F. jacksoni, 133, 184
pondicerianus, F., 109, no, 117, 122, 167,
169, 179, 196
pondicerianus, F. p., 167, 196
prepositus, F. pondicerianus, 179
proximus, F. africanus, 177
psilolaemus, F., no, 117, 121, 122, 152-160,
169, 177, 192
psilolaemus, F. p., 153, 192
punctulata, F. afer, 175
rovuma, F. sephaena, no, 150, 177, 190
ruahdae, F. coqui, 162, 164, 178, 194
ruandae, F. coqui, 176, 178
ruandae, F. nobilis, 176
rufopictus, F., no, 113, 116, 125-131, 169,
170, 182
sarudnyi, F. francolinus, 175
schlegelii, F., no, 114, 117, 118, 160-165,
1 66, 169, 170, 179, 194
schoanus, F. sephaena, 177
schubotzi, F. lathami, 166, 196
schuetti, F. squamatus, 137, 138, 139, 176,
186
sephaena, F., no, in, 113, 117, 118, 121,
123, 148-152, 167, 169, 177, 190
sephaena, F. s., 150, 152, 177, 190
sequestris, F. shelleyi, 177
sharpii, F. clappertoni, 144, 147, 176, 188
shelleyi, F., no, in, 113, 115, 117, 118, 121,
122, 152-160, 168, 169, 173, 177, 192
shelleyi, F. s., in, 113, 122, 153, 154, 155,
159, 160, 168, 177, 192
somaliensis, F. sephaena, 177
spilogaster, F. sephaena, 150, 177, 190
spinetorum, F. coqui, 114, 161, 162, 165, 194
squamatus, F., no, in, 116, 118, 121, 136-
140, 1 66, 1 68, 169, 176, 1 86
squamatus, F. s., 137, 138, 139, 186
stantoni, F. levaillantoides, 155, 178
streptophorus, F., 110, 114, 117, 120, 148-
152, 167, 169, 170, 171, 172, 190
stresemanni, F. levaillantoides, 178
stuhlmanni, F. coqui, 178
swainsonii, F., no, 116, 121, 122, 125-131,
169, 175, 182
swainsonii, F. s., 127, 129, 130, 182
swierstrai, F., no, 116, 131-136, 169, 176,
184
swynnertoni, F. afer, 127, 128, 131, 175, 182
testis, F. clappertoni, 176
tetraoninus, F. squamatus, 176
thamnobium, F. natalensis, 176
theresae, F. psilolaemus, 153, 154, 192
thikae, F. coqui, 161, 162, 164, 194
thompsoni, F. sephaena, 177
thornei, F. bicalcaratus, 144, 188
titar, F. pondicerianus, 179
tokora, F. leucoscepus. 175
tornowi, F. afer, 175
trothae, F. shelleyi, 177
tschadensis, F., 141, 176, 188
ugandensis, F., 141, 176, 188
uluensis, F. shelleyi, in, 154, 155, 159, 160,
168, 192
usambarae, F. squamatus, 137, 139, 186
uzungwensis, F. squamatus, 137, 139, 186
vernayi, F. coqui, 163, 194
watsoni, F. pictus, 124, 180
wattii, F. levaillantoides, 178
wellsi, F. pintadeanus, 175
whitei, F. squamatus, 176
whytei, F. shelleyi, in, 113, 122, 154, 155,
157, 159, 160, 192
zambesiae, F. sephaena, 150, 151, 152, 177,
190
zappeyi, F. squamatus, 176
zuluensis, F, sephaena, 177
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED,
BARTHOLOMEW PRESS, DORKING
TAXONOMY AND BIOLOGY
OF THE GENUS LEBETJ7S
(TELEOSTEI-GOBIOIDEA)
P. J. MILLER
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 10 No. 3
LONDON: 1963
TAXONOMY AND BIOLOGY OF THE GENUS
LEBETUS (TELEOSTEI-GOBIOIDEA)
BY
P. J. MILLER
(Department of Zoology, The University, Glasgow, W.2)
Pp. 205-256 ; Plate ; 21 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 10 No. 3
LONDON : 1963
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(NATURAL HISTORY), instituted in 1949, is
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TAXONOMY AND BIOLOGY OF THE GENUS
LEBETUS (TELEOSTEI-GOBIOIDEA)
By P. J. MILLER
ABSTRACT
The marine gobioid genus Lebetus Winther 1877 is redefined and previous division into two
species — orca Collett 1874 and scorpioides Collett 1874 — shown to be based on sexual dimorphism
in one species, by page priority L. orca. A redescription of the species is provided including
details of sensory papillae and osteology. L. orca is eastern Atlantic boreal in distribution,
occurring as a small predator on coarser grounds down to 375 m. Specialization of the male
genitalia and sexual dimorphism are described and the breeding season provisionally regarded
as from March to August. The skeleton of Lebetus is typically gobiid, and evidence is presented
for a closer relationship to the genus Buenia Iljin as represented by B. Jeffrey sii (Giinther) than
to the Mediterranean Odondebuenia De Buen and Cabotichthys Whitley with which Lebetus had
been previously grouped.
INTRODUCTION
THE genus Lebetus was founded by Winther (1877) to contain two species of gobies
dredged by G. O. Sars on the coast of Norway and described by Collett (1874) as
Gobius orca and G. scorpioides. These species have subsequently been regarded as
among the rarest of European Atlantic fishes and, until recent years, the numbers of
demersal individuals known to science have been relatively few. In the last fifteen
years, protracted dredging surveys off the Isle of Man (Jones, 1951 ; Hartnoll, 1961)
have yielded many examples. Their small size and occurrence on rough grounds makes
collecting difficult ; in Manx waters, most specimens have been obtained by means of
a scallop dredge lined with shrimp netting. Relative infrequency of capture is
probably due to inadequacies of the fishing gear rather than to actual rarity, and
occasional hauls providing up to five individuals suggest the presence of appreciable
numbers in the area. Other long term faunistic investigations within the range of the
genus have produced more or less sizeable collections of both demersal (Grieg, 1913 ;
Le Danois, 1913) and planktonic stages (Petersen, 1919 ; Johansen, 1925 ; Russell,
1926-1940).
My interest in these fishes was aroused firstly by the difficulty experienced in
separating the two species. After endeavouring to use the diagnoses of several authors,
it was found possible to achieve this only on the basis of coloration. Then, while
examining the testes of various gobies, it was noticed that no female L. orca were
present in a collection of twenty two dissected individuals, and only immature males
among twenty eight L. scorpioides. The possibility that the two so-called species
were in fact based on sexual dimorphism in a single species indicated the need for
taxonomic revision. Although such a view was put forward by Smitt (1900), this
was not accompanied by any evidence and appears to have been overlooked by later
authors. Dissection of the extensive material from the Irish Sea presented oppor-
tunities for settling the position of Lebetus among the gobioid families by osteological
ZOOL. 10, 3 i4§
208 P. J. MILLER
study, and also for investigating various aspects of diet and reproduction. A pre-
liminary account of some of this work has already been published (Miller, 19616).
MATERIAL AND METHODS
The following specimens have been available for examination :
Isle of Man : 33 $<$, standard lengths 15 -75-30-0 mm., including BMNH 1961 . 10 . 17 .
i. 3» 5» an(i one m PEM, and 27 ?, 14-5-29-0 mm., inc. BMNH 1961 . 10 . 17 . 2, 4,
and one in PEM.
Other British localities : 3 £$, 12-0-30-0 mm., BMNH 88 . 3 . 22 . 9-10 part and BMNH
1903.4.14.4-7 part, and 2 $$, 10-5 and 17-0 mm., BMNH 1903.4.14.4-7 part.
Scandinavia: 13 <$<$, 13-5-26-0 mm.,ZMO J3999 (type of Gobius orca), J40OO, J4OOI,
J4023 ; ZMB 771, 1887, 1893, 1966, 4174, 4175, 5294 ; ZMC 89, 91 ; and 4 ??,
21-75-29-0 mm., ZMO J402O, J4O2I (types of G. scorpioides) ; ZMB 536, 2009.
Abbreviations for museums are as follows :
BMNH British Museum (Natural History).
PEM Port Erin Marine Biological Station Museum.
ZMB Zoologisk Museum, Universitetet i Bergen.
ZMC Universitetets Zoologisk Museum, Copenhagen.
ZMO Zoologisk Museum, Universitetet i Oslo.
The specimens were viewed by means of low power binocular and dissecting micro-
scopes. With suitable illumination the sensory papillae were readily seen without
staining after careful removal of mucus from the head and body surfaces. The skeleton
was investigated by dissection of alizarin stained whole-mount preparations.
THE GENUS LEBETUS WINTHER 1877
Lebetus Winther, 1877: 49 (orthotype Gobius scorpioides Collett 1874); Smitt, 1900: 554;
Jordan, 1919 : 392 ; 1923 : 225 ; Duncker, 1928 : 140 ; Iljin, 1930 : 55 ; De Buen, I93oa :
123 ; 19306 : 5, 21 ; Koumans, 1931 : 43, 46 ; Whitley, 1931 : 155 ; De Buen, 1931 : 50, 54,
61 ; 1940 : 3 ; 1951 : 56, 57 ; Norman, MS : 413.
Lebistes Jordan, 1920 : 487 ; 1923 : 226 (non Filippi, 1862 : 69).
Butigobius Whitley, 1930 : 123.
Body moderately elongate, subcylindrical, with 25-29 ctenoid scales in lateral
series. Head, nape, and back naked to origin of second dorsal fin, lacking skin folds
or barbels. Postorbital length not more than half length of head. Anterior nostril a
short tube. Opercle without scales. Preopercle and lower jaw unarmed. Branchi-
ostegous membranes attached to sides of isthmus, but not fused across isthmus. Jaws
oblique, subequal, maxillae ensheathed and not elongate. Teeth of both jaws in
several rows, simple, erect. Tongue truncate to very weakly emarginate, free from
floor of mouth anteriorly. Pelvic fins united along entire length, without anterior
membrane (frenum) and with fourth branched ray somewhat produced. Pectoral
girdle lacking flaps on anterior edge ; uppermost rays of pectoral fin not free from
membrane. Dorsal fins separate, first dorsal with 6 rays, not pungent. Caudal fin
rounded, not much longer than head.
Cephalic canals absent. Sensory papillae prominent but reduced in number. Two
pairs of interorbital papillae (/>). Cheek with only suborbital row a, a short longi-
TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 209
tudinal row c, and an intermediate papilla ; row d without posterior longitudinal
section. Anterior dorsal (occipital) series (excluding h) in two groups of a few papillae.
Opercular transverse row ot in two sections. Preoperculo-mandibular series in
several parts.
In compiling his work on the genera of fishes, Jordan (1919, 1920, 1923) includes,
together with Lebetus Winther 1877, the genus Lebistes Smitt 1899. Since Jordan
gives " L. scorpioides Smitt " as the orthotype of this genus, the latter, as noted by
Koumans (1931), is clearly an erroneous reference to Lebetus for which Smitt (actually
1900, not 1899) indicated Gobius scorpioides Collett as the type. The generic name
Lebistes is preoccupied by Lebistes Filippi 1862 among the cyprinodont teleosts and,
to replace this name within Jordan's list of gobiid genera, Whitley (1930) proposed
Butigobius without consulting Smitt 's paper. In a later account, Whitley (1931)
unravels this confused story.
THE SPECIES OF THE GENUS
The present section embodies a review of the various characters proposed by
earlier authors as being of value in the diagnosis of L. orca and L. scorpioides.
Coloration. The two species are reputed to differ in both body and fin coloration.
The body of Gobius orca was described by Collett (1874, 18750) as uniformly greyish
except for darker areas between the second dorsal and anal fins, and at the root of
the caudal fin, while that of G. scorpioides was said to possess four or five dark brown
vertical bands across the sides. After examination of a living scorpioides, Winther
(1877) remarked on the colourless caudal peduncle, and both Holt & Byrne (1903)
and Le Danois (1913) emphasized the importance of this pale band, sharply demar-
cated anteriorly, as a means for the identification of L. scorpioides. In the original
description, Collett (1874) mentioned that the second dorsal fin of G. orca had an
indistinct white band in its outer half whereas the same fin in G. scorpioides showed
only ill-defined banding. Alternate dark brown and snow white bands were found
across the second dorsal fin of a later specimen of G. orca (Collett, 1885). Holt &
Byrne (1903) regarded a superior black band to this fin as characteristic of G. orca
and in their text referred to a row of white spots immediately below the dark peri-
phery. These authors described and illustrated the second dorsal fin in G. scorpioides
as banded with red, the markings in some cases being flanked by dark edging.
With the collection of Lebetus amassed for this work, it has been possible to segre-
gate the individuals into two groups on the basis of differences in the coloration of the
second dorsal fin. In the first of these, termed the orca group, the second dorsal fin
has a thick black superior edge, below which occur white areas and wide oblique
bands, yellow or ochre in life but grey in preserved material. Within the scorpioides
group, the black edge to the second dorsal fin is present but much thinner and less
intense than in the orca examples. The rest of the fin bears merely four narrow double
lines of melanophores, enclosing in the living fish thin red or ferruginous striae, and
separated by hyaline fin membrane. Among living scorpioides, madder brown lateral
marks are pronounced on the body which, in orca fishes, is more uniform grey or.
yellowish grey although comparable markings may be quite prominent. A pale band
on the caudal peduncle is more clearly defined in the scorpioides group, especially
2io P. J. MILLER
when preserved, but living orca show this feature to a noticeable degree (cf. PI. I and
Text-fig. 3) and it is shown in Collett's illustrations of Gobius orca (i875a, 1885).
TABLE I. — Relation of Coloration Groups to Sex and Maturity Stage
No. of males
Mature and No. of females
Group developing Immature all stages
(a) Dissected
orca 21 i
scorpioides . 6 22
(b) Not dissected
orca . . 16
scorpioides . 5 n
The relation of these two colour patterns to sex and to gonad maturation is indicated
in Table la, based on fifty Manx specimens whose sex and sexual development were
confirmed by dissection. The sex and approximate maturity stage in a further thirty
two British and Scandinavian examples have been determined by inspection of the
genital papilla (Text-fig. 17), and these results are given in Table Ib. It is apparent
that mature or developing male scorpioides do not occur, and that female orca are
unknown at any stage of maturation. This distribution strongly implies that the
orca coloration is the male livery of a species whose females and immature males bear
the markings of scorpioides. Incipient development of the orca pattern in the second
dorsal fin can, in fact, be noted among some of the immature males in the scorpioides
group, which usually exhibit a pronounced black spot on the distal part of the first
two interradial membranes of the second dorsal fin (see Le Danois, 1910, fig. 4).
The above hypothesis is further supported by the range in standard length found in
the three categories shown in Table II. The examples of orca are from 18-0-30-0 mm.
TABLE II. — Standard Length Frequency in Coloration Groups, Expressed in 2 mm.
Length Groups
Standard length
(
Coloration group
orca .
cJ scorpioides
9 scorpioides
in length while male scorpioides do not exceed 23-0 mm. Taning (1940) has reported
juvenile Lebetus orca of 5-15 mm., but relied for the identification on meristic
characters whose validity is doubtful. Female scorpioides attain about the same
maximum length as that for orca.
The first dorsal fin in the orca individuals is a uniform yellowish grey edged with
white, but in scorpioides is usually banded with a dark spot on the interradial mem-
brane between the fifth and sixth rays. Confluence of these bands frequently occurs,
and the spot may not be very distinct. This scorpioides group pattern is stated by
10
12 14
16
18
20
22
24
26
25
1
30
i
i
IO
16
7
i
2
2 3
3
2
I
I
I
2
i
2
5
9
5
7
TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS an
Holt & Byrne (1903) to be found only in the breeding male of " Gobius scorpioides ".
Fage (1918) regarded the presence of a black spot on the first dorsal fin of a 12-5 mm.
example from the Dana collections as a sign of precocious development of supposed
male coloration. However, in the material under consideration, the spot is well
represented in adult females and among male Lebetus occurs only in the immature
where it is not uncommonly absent. The single female G. orca mentioned in the
literature was taken by Patience (1906) in the Firth of Clyde. This specimen has
not been traced. The diagnosis of G. orca given by Patience was not derived from his
own examples but seems to be a translation from Collett (1896) and could apply to
individuals of either the orca or scorpioides group.
TABLE III. — Meristic Characters Cited for Lebetus orca and L. scorpioides
No. of fin rays
First
Second
•\
Verte-
dorsal
dorsal
Anal
Pectoral
Caudal
brae
L. orca1
VI
ii
10
18
3/13/3
VI
ii
9
17
3/13/3
VII
IO-II
9
17
3/12-13/3
VII
I/9-io
1/8
17
12-13
VI-VII
I/9-io
1/8
17
x/13/x
VII
IO-II
9
17
3/12-13/3
VI-VII
9-1 1
9-10
VI-VII
I/9-io
1 8 (sic)
17-18
12-13
VI-VII
I/9-io
1/8-9
VI-VII
10-12
8-10
18-20
7-8/1 1/6-7
28-29
L.
scorpioides2' 3
VI
9
8
20 (18)
6/12/6
VI
9
8
1 8-20
6/12/6
VI
9
8
c. 18
12
VI
1/8
1/7
18-20
12
VI
1/8
1/7
18-20
X/I2/X
VI
VI
VI
VI
VI
V-VI
VI
9
9
1/9
1/8-9
8-9
1/7-8
1/8-9
8
7-8
1/6
1/6-9
6-8
1/6-7
1/6-7
18-20
18-20
18-20
X/I2/X
12
X/I2/X
No. of
Scales
in 1.1.
28
28
24
25
c. 24
25-28
25-26
24-28
28
28
30
28-30
c. 28-30
28-30
28
28
27-28
28
25-26
25-26
26-30
28-30
Authority
Collett, 1874
Collett, 18750
Collett, 1885
Lilljeborg, 1884
Smitt, 1892
Collett, 1896
Holt & Byrne,
1903
Duncker, 1928
De Buen, 19300, b
Taning, 1940
Collett, 1874
Collett, 18750
Winther, 1877
Lilljeborg, 1884
Smitt, 1892
Holt & Byrne, 1903
Le Danois, 1910
Le Danois, 1913
Fage, 1918
Petersen, 1919
Duncker, 1928
De Buen, I93oa,
b, 1932
1 Scales in 1.1. c. 26-c. 28 (Grieg, 1913). z Scales in 1.1. 26, T)l V (Collett, 1902). 3 Scales in 1.1.
28 (Grieg, 1913)-
Meristic characters. Radial formulae, vertebral and scale counts provided by
various authors for the two species are shown in Table III. Although Collett origin-
ally ascribed 6 elements to the first dorsal fin of both species, he later changed this to
212 P. J. MILLER
7 for G. orca. Lilljeborg (1884) and Smitt (1892) employed this supposed disparity in
keys to Scandinavian gobies. L. orca is also regarded as having rather more articu-
lated rays in the second dorsal and anal fins than L. scorpioides, and slight differences
are indicated in pectoral and caudal fin ray counts. The number of scales along the
lateral midline is reputed to be greater in L. scorpioides than in L. orca.
For the orca and scorpioides groups of the present material the meristic values in
Table IV have been obtained. Methods of counting are described in the systematic
section below. To minimise the effect of regional variation, results for British and
Scandinavian collections have been kept separate. Mean values are given for the
more numerous British material.
TABLE IV. — Meristic Characters of Coloration Groups : Number of Observations
(Highest in Bold Type) Against Value. M — Mean Value for British Material
Origin of specimens
Coloration group
orca
scorpioides
orca
scorpioides
orca
scorpioides
orca
scorpioides
orca
scor
orca
scorpioides
orca
scorpioides
British Isles
5 6
26
34
First dorsal fin rays
Scandinavia
6 7
7 I
6
Second dorsal fin articulated rays
6 7 8 9 10 ii M 8 9 10 ii
i 18 7 9-23 531
i 2 4 21 6 8-85 2 4 i
Anal fin articulated rays
5 6 7 8 9 M. 56789
i 19 6 7 -19 54
32 22 7 6-97 i 4i
Pectoral fin rays
15 16 17 18 19 20 21 M 17 18 19 20 21
2 15 20 15 18-92 742
i 4 7 9 30 10 i 18-56 4 2
Caudal fin branched rays
8 9 10 ii M 8 9 10 ii
24 i 9 -04 5
2 31 i 8-97 2
Scales in lateral series
25 26 27 28 29 M 25 26 27 28 29
2 18 10 6 2 26-68 i 3 2 i
i 17 17 6 i 26-74 i i 2
Vertebrae including urostyle (Manx specimens only)
26 27 28 29 M
2 13 i 27-94
i 3 17 27-76
TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS
213
Except for pectoral ray and scale counts in the Scandinavian fishes, where the
number of observations is small, the most common value for each meristic character
is the same in the two coloration groups and mean values are only slightly different.
The wider range and somewhat lower means for fin ray counts in scorpioides are due
17-
< 15-
13-
Original standard Present
First
Second
Pect-
Scales
Sex
ZMO No. .
+ Caudal fin
standard
Dorsal
dorsal
Anal
oral
in 1.1.
6*
J 3999
26-0 + 6-0
25-0
d.
I/IG
1/8
d.
26-27
?
J 4020
total 28-0
21-5
VI
1/9
1/7
d.
d.
?
J 4°21
30-0 + 7-0
29-0
VI
1/9
1/7
20
27-28
TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 219
GENERAL DESCRIPTION. Body subcylindrical, laterally compressed towards
caudal fin ; in standard length, depth at origin of pelvic fin 5-1-6-65 (mean 5-65), at
origin of anal fin 5-7-7-5 (6-53), of caudal peduncle before origin of caudal fin 8-5-12-3
(10-12). Head rounded ; in standard length, horizontal length (snout to upper origin
of opercle) 3-2-4-1 (371), maximum width (between upper origins of opercles)
5'25-5 '95 (5 '68). Horizontal distance from tip of snout to origin of first dorsal fin
2.7-3-05 (2-9), to origin of second dorsal fin 1-7-1-9 (1-81), to anus 1-75-1-95 (1-86),
to origin of anal fin 1-6-1-75 (1-69), to origin of pelvic fin 3-2-4-0 (3-47), all in standard
length. Caudal peduncle (end of anal fin base to origin of caudal fin) horizontal
length 4-15-5-1 (4-71) in standard length, depth 1-8-2-35 (2*°7) m own length. In
head length, snout 3-8-5-6 (4-7), eye 2-8-3-4 (3-0), postorbital length 2-0-2-7 (2*34),
cheek 3-8-4-85 (4-34) ; eye and cheek direct, rest horizontal measurement. Inter-
orbital minimum width (bony) 4-3-7-4 (5-5) in eye length. Snout less than diameter
of eye, with moderately sloping profile ; nostrils adjacent, anterior on short tube,
not projecting over upper lip and with entire distal rim lacking local prolongation ;
posterior nostril also tubular but shorter. Eyes large, close together. Postorbital
region never exceeding half head length ; profile more or less flat, without dermal
ridges. Upper jaw relatively wide, about equal to preorbital area. Mouth oblique,
lower jaw at most only slightly in advance of upper ; maxilla ends posteriorly under
anterior quarter of eye. Lower jaw unarmed, without barbels. Cheek smooth, without
dermal ridges. Preopercle unarmed. Branchiostegous membrane attached to entire
lateral margin of isthmus from immediately anterior to lower edge of pectoral lobe and
without transverse fusion across isthmus (Text-fig. 5). Teeth simple, caniniform, erect
in both jaws. Dentition of jaws an outer row of larger teeth with narrow band of two
to four rows of smaller teeth. Pharyngeal teeth relatively large, recurved, in two
superior and one inferior median patch (see description of skeleton). Tongue more or
less truncate, with anterior edge rounded laterally ; at most pronounced, a very
slight median emargination (Text-fig. 6s). Gill rakers small protuberances, not
spinous, about 6-8 on first arch. Sagittal otolith rectangular, anterior and posterior
edges slightly concave, inferior edge slightly convex, superior edge with angular
projection near posterior end ; outer surface convex, more flattened in middle, inner
surface convex, with furrow in middle and shallower groove to posterior edge (Text-
fig. 6A).
FINS. In the following counts, spinous elements are represented by large roman,
articulated rays by arabic numerals. The terminal bifid ray in the second dorsal
and anal fins is counted as one element. Only in the caudal and pelvic fins are articu-
lated rays branched. The range of values is given in parentheses after the most
common number, frequency distribution being shown in Table VII.
First dorsal fin VI (VI-VII) ; second dorsal fin 1/9 (6-n) ; anal fin 1/7 (5-8) ;
caudal fin (branched rays) 9 (8-10) ; pectoral fin 19 (15-21), pelvic fin 1/5 -f- 1/5.
In standard length, first dorsal base 7-95-8-95 (8-24), second dorsal base 3-75-4-55
(4-24), anal base 4-85-5-8 (5-4), longest caudal ray 3-2-4-45 (for standard lengths up
to 20-0 mm., mean = 3-55 ; more than 20-0 mm., mean — 3-92), longest pectoral
ray 3-7-4-45 (4-0), pelvic origin to tip of fifth pelvic articulated ray 3-85-4-55 (S.L.
up to 25 mm., mean = 4-05 ; more than 25-0 mm., mean = 4-2). First dorsal fin
P. J. MILLER
TABLE VII. — M eristic Characters of Lebetus orca : Number of Observations
against Value
Locality
British Isles
Scandinavia
British Isles
Scandinavia
British Isles
Scandinavia
British Isles
Scandinavia
British Isles
Scandinavia
British Isles
Scandinavia
Isle of Man
First dorsal fin rays
5 6 7
60
Second dorsal fin articulated rays
6 7 8 9 10 ii
i 2 5 39 13
2941
Anal fin articulated rays
567*9
3 3 4i 13
i 9 5
Pectoral fin rays
75 16 I^ 18 19 20 21
i 4 9 24 50 25 i
784
Caudal fin branched rays
8 9 10 ii
2 55 2
7
Scales in lateral series
25 26 27 28 29
3 35 27 12 3
1433
Vertebrae including urostyle
26 2j 28 29
i 5 30 i
TABLE VIII. — Dorsal Fin Ray Lengths Expressed in Standard Length for Mature and
Immature Lebetus orca
Mature • OOO 0
6
• • •
• too
• • • • «o •
• ®o • o
• • •• 0
o
5
•• 0> 0>
•
•O* COD O°
0 0
» »o * o o o
*
4-
0*0000
•
• • • o
Q
o
3-
o
^ o o
2
* 0 • O
0
•
o
•
^ 2-
1"
o
.c 1 •
"5)
c
_l
X
0
oe
FIRST DORSAL
FIRST DORSAL
.E 8-
u_
'" I
IV J
7-
• ^
. \I
• •• o> o
• .1 •
6-
• *o
• ••
• • O O ° 00
t o
•o o o o
• 0
5-
o o o
Q
• • 0
000
• CD
O
O CB O
OO O O
O
o o
o o
o
o • o
O oq
o o e o
o
O OCX OOO
p
1 — | - 1 — | - 1
15
1 — I - 1 — I
20
— 1 — I — 1 - 1 — 1 — 1
25 30
—I — I — I — I — I — I —
15 20
25 30
Standard Length (mm)
FIG. 20. Relation between standard length and length of last two rays of first dorsal fin
(V, VI) and first two rays of second dorsal fin (I, i) in L. orca. Symbols as in Fig. 19.
242 P. J. MILLER
eight diverse genera of gobies. The group is characterized by several peculiarities.
Among these are the large non-osseous area between the symplectic, quadrate, and
preoperculum ; the arrangement of the branchiostegals on the ceratohyal ; the absence
of the parietal bones ; and the presence of a splintlike radial above and below the
hypurals in the caudal skeleton. In all these features, the genus Lebetus appears as a
typical gobioid fish, but there has been some doubt about the position of Lebetus among
the families of this suborder. Of these about six are now recognized (Koumans, 1953 ;
Gosline, 1955 ; Norman, MS.). The Rhyacichthyidae, Taenioididae, Kraemeriidae, and
Microdesmidae are highly aberrant gobioids and need not be further dealt with in this
discussion. The two largest and most important families are the Eleotridae and
Gobiidae. Skeletal differences between these have been summarized by Regan (1911),
and the diagnostic value of certain features reviewed by Gosline (1955). Nevertheless,
in referring gobies to one or other of these families, it has been customary to use the
external form of the pelvic fins as the chief guide in the usual absence of osteological
data. According to Regan (1911) and other authors, the Eleotridae are characterized
by the pelvic fins being separate, and the Gobiidae, when these fins are developed, by
their union to produce a simple cup-shaped disc. However, various stages in the
union or separation of the pelvic fins have been recognized among gobioid genera and
the importance of this feature as a criterion of family position has recently been
questioned, especially by Bohlke & Robins (1960^, b) who have found eleotrid, gobiid,
and intermediate types of pelvic fins in a single genus (Coryphopterus Gill). In such
cases as these, determination of systematic position within the suborder requires
investigation of the skeleton.
When erecting the genus Lebetus, Winther (1877) emphasized that the anterior
membrane of the pelvic disc was missing, although the two fins were joined along their
entire length in the midline. At first accepting (1885) the new genus, Collett (1896)
was later of the opinion that intermediate stages in reduction of the anterior mem-
brane could be found between the fully developed condition in Gobius and complete
loss as reported for Lebetus, and doubted the validity of separating the latter from
Gobius. On the other hand, Duncker (1928) found that the two pelvic fins were
completely separated in Lebetus but stressed the desirability for re-examination of
these fins in better preserved material in order to decide whether Lebetus belonged
to the Eleotridae or the Gobiidae. De Buen (19306) associated Lebetus with two
Mediterranean genera, one of which had previously been regarded as an eleotrid
because of its almost completely separated pelvic fins, and placed this " Lebetus "
group in the subfamily Gobiinae of the Gobiidae. Koumans (1931) similarly accepted
Lebetus as a gobiid genus. In the present work, it has been confirmed that, in the least
damaged examples, the pelvic fins of Lebetus are completely united between their
origins and the distal extremity of the fifth ray of each fin, but are entirely without
trace of an anterior membrane to complete the disc. To settle the question of family
position with greater certainty, it became obvious that examination of the skeleton
was necessary, and the results of this study reveal that Lebetus has the characters of
a typical member of the Gobiidae. These include a T-shaped head to the palatine,
loss of the mesopterygoid from the palatopterygoid arch, and absence of a hyper-
coracoid in the pectoral girdle, where the laminar radials are inserted on the cleithrum
TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 243
and the lowest also related to the hypocoracoid. In the possession of a spatulate
glossohyal, five branchiostegous rays, a reduced metapterygoid, and loss of the
opisthotic, Lebetus also reaches a higher level of specialization than that described
among the eleotrids (Regan, 1911 ; Takagi, 1950 ; Gosline, 1955).
Within the Gobiidae, the affinities of Lebetus undoubtedly lie with the subfamily
Gobiinae, since the genus exhibits none of the more extreme modifications of the
Sicydinae, Periophthalminae, Apocrypteinae, Tridentigerinae, Gobiodontinae, or
Benthophilinae (Koumans, 1953 ; Norman, MS.). In the absence of an anterior
membrane to the pelvic disc, Lebetus resembles a number of small Indo-Pacific
genera i.e. Herreogobius Koumans, Quisquilius Jordan & Everman, Fusigobius
Whitley, Amblyogobius Bleeker, and Zonogobius Bleeker (Koumans, 1953 ; Gosline,
1959) . With head and anterior part of body naked, elongate rays in the first dorsal
fin, 22-30 scales in lateral series, and radial formula of D2 1/8-9, A I/7~8 (Koumans,
I953)> Zonogobius appears to be closest of these to Lebetus. Differences comprise
the larger gill openings, and more laterally compressed head in Zonogobius. However,
it is impossible to trace relationships between European and exotic genera, when the
arrangement of the lateral line system in so many of the latter, including Zonogobius,
remains to be adequately described and figured. As shown in the normal illustration
of Zonogobius corallinus sp. nov. (Mozambique) by Smith (1959, Fig. 29), the dis-
tribution of sensory papillae on cheek and postorbital regions differs from that of
Lebetus in the complete longitudinal rows and more numerous papillae. Koumans
(1931) associated Lebetus with the genus Coryphopterus Gill ; as defined by Koumans,
the latter included European Pomatoschistus species together with the genotype, C.
glaucofraenum Gill, from the tropical Western Atlantic. In a revision of Coryphop-
terus sensu stricto, Bohlke & Robins (19606) provide details of the cephalic lateral
line system and other characters which do not point to any close affinity with Lebetus,
in spite of the fact that two of their new Western Atlantic species (C. alloides and C.
dicrus) are without the anterior pelvic membrane.
In the European fauna there are two Mediterranean genera of gobies lacking
anterior membranes to the pelvic fins. These are Odondebuenia and Cabotia, both
introduced by De Buen (19306). Since Cabotia is preoccupied among the Lepidoptera,
De Buen (i5th June, 1940) suggested Fagea as a replacement but was preceded in this
by Whitley (May, 1940) who introduced Cabotichthys. The genus Odondebuenia
contains two species which were originally (1907) placed among the Eleotridae as
Eleotris balearicus Pellegrin & Fage and E. pruvoti Fage. Odondebuenia is character-
ized by pelvic fins united only at their bases, naked nape and throat, unusual gill
rakers (Fage, 1907, figs. 3 and 9), modified scales at the origin of the caudal fin, and
meristic characters of D2 1/9-11, A I/8-io, and Sc. 1.1. 24-32. Both species of
Odondebuenia show sexual dimorphism in length of first dorsal fin rays (Fage, 1918 ;
De Buen, 19306). The second of these Mediterranean genera, Cabotichthys, has the
pelvics joined together for about half their length, nape and throat fully scaled, scales
of the caudal peduncle not modified, and fin ray and scale counts of D2 1/14, A 1/13,
and Sc. 1.1.50. The genus is founded on one species, C. schmidti (De Buen) known
only from the single type specimen (De Buen, 19306). De Buen (1931) included
Odondebuenia, Cabotichthys and Lebetus in his " Lebetus " group of genera, the principal
244 P. J- MILLER
character of which is the absence of the anterior pelvic membrane. Similarities in habi-
tat also link these fishes, all three being obtained on coarse grounds where calcareous
algae are noticeable constituents of the bottom deposits, and none exceed 50 mm. in
length.
On the face of the diagnoses given above, Lebetus would appear to be fairly closely
related to Odondebuenia except for the occurrence in the latter of modified caudal
scales and gill rakers, and the greater separation of the pelvic fins, while Cabotichthys
stands quite distant from both in several respects. The lateral line system in Odonde-
buenia and Cabotichthys has fortunately been investigated by De Buen (19306, 1940)
and that of Lebetus is described above. Comparison of these accounts for the three
genera reveals the heterogenous nature of their grouping. The patterns of sensory
papillae and extent of the cephalic canals in Cabotichthys point to an intimate connec-
tion with the genus Gobius, as defined by De Buen (19300, 1931), and, apart from the
reputed form of the pelvics, other characters of this fish do not warrant any generic
separation from Gobius. My own experience with dredged and trawled examples of
Lebetus and other gobies is that the pelvic membranes are very susceptible to damage,
and the pelvics of Cabotichthys, as figured by De Buen (19306, fig. 7) are reminiscent
of such a condition. It may well prove that C. schmidti is no more than a damaged
specimen of a Gobius species.
The state of the pelvics in the Odondebuenia species seems well established from a
number of specimens. In the arrangement of sensory papillae, this genus differs
markedly from Lebetus in the greater number of papillae, and their distribution in
well marked transverse and longitudinal rows on the cheek approaches that in Gobius,
which Odondebuenia also resembles in the possession of a similar cephalic canal
system. The morphological attributes common to both Odondebuenia and Lebetus
are not of a very highly specialized nature and do not outweigh the considerable
disparity in patterns of sensory papillae. Modification of scales at the base of the
caudal fin involves pronounced elongation of all the ctenii in 0. pruvoti (Fage, 1907,
fig. 7) or merely the lateral ctenii in 0. balearica (Fage, 1907, fig. n ; De Buen, 19306,
fig. 2). Although not seen in Lebetus, it is of interest to note the occurrence of this
peculiarity in the new gobioid genus Various, recently described from the West
Indies by Robins & Bohlke (1961) and also with separated pelvic fins. These authors
commented on the existence of comparable scales in the Californian eleotrid Chriolepis
Gilbert and the gobiid Garmannia Jordan, which is represented on both sides of
Central America and in the Caribbean. It is not proposed at this point to deal
further with the relationships of Odondebuenia, except to call attention to the need
for an osteological investigation of this genus.
Loss of the anterior membrane from the pelvic disc has probably occurred on a
number of different evolutionary lines among the Gobiidae and, as in the case of
Odondebuenia and Lebetus, need not be a sure indication of close phyletic association.
In geographical distribution Lebetus appears to be confined to the eastern Atlantic
boreal region and may well have evolved there. It is in this area, therefore, that the
closest relatives of the genus may be sought. Eight other gobioid genera are recorded
from the eastern North Atlantic (De Buen, 1931) and all of these have a pelvic disc
complete with anterior membrane. The two pelagic forms Aphya Risso and Crystal-
TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 245
logobius Gill may be excluded from consideration on the grounds of extreme specializa-
tion. Among the demersal gobies, the genera Deltentostem Gill, Pomatoschistus Gill,
Chaparrudo De Buen, and Gobius L. exhibit various features of anatomy and lateral
line organization (see De Buen, 1930^, 1931) which do not suggest any near relation-
ship with Lebetus. The remaining genera, Lesueurigobius Whitley and Buenia Iljin,
deserve more attention. The first of these, represented by Lesueurigobius friesii
friesii (Collett), has a number of characters in common with Lebetus. Thus both
genera are entirely without cephalic canals and in disposition of sensory papillae
show a posterior extension of row a behind the eye, papillae in the oculoscapular
furrow (row u), interorbital papillae, a high posterior termination for row i, and reduc-
tion of rows m, n, and o (Sanzo, 1911, pi. 9, figs. 4, 5, as Gobius macrolepis ; De Buen,
1923, figs. 21, 22). However, the great abundance of papillae in Lesueurigobius
contrasts with their relative scarcity in Lebetus, where several rows (g, x, z, c2, b, d)
found in Lesueurigobius are little if at all developed, and others (i, e, ot, c-cp] inter-
rupted. Except for reduction in rows m, n, and o, the resemblances listed may be
accounted for by independent action of a similar evolutionary process in the past
history of each genus. This was the replacement of existing cephalic canals by rows
of sensory papillae. Considerable anatomical differences in size, habit, squamation,
radial formulae, and coloration (Holt & Byrne, 1903 ; Duncker, 1928) would also
suggest similarities are due to convergence, and that any affinity is relatively distant.
No closer relationship can be demonstrated with Lesueurigobius sanzoi (De Buen) or
L. lesueuri (Risso) from the Mediterranean (De Buen, 1923).
The only genus now remaining is Buenia, the Atlantic species of which is B. Jeffrey sii
(Giinther), and it is with this form that Lebetus appears at present to be most closely
connected. Lacking scales on the head and predorsal regions, and with meristic
characters of D2 1/8-9, A I/y-8, and Sc. 1.1.25-30 (Duncker, 1928 ; personal
observation), B. Jeffrey sii displays a resemblance to Lebetus which is also evident in
the arrangement of the lateral line system (Text-fig. 21). On the cheek absence of
transverse rows of papillae, and the short row of large papillae with an intermediate
papilla between this and row a, recall the conditions in Lebetus. The lateral preorbital
rows c1 and c2 are identical in both genera. On the dorsum of the head there is a
correspondence in the reduced number and the arrangement of the papillae. Lebetus
differs from B. Jeffrey sii in the disappearance of cephalic canals and their replacement
by papillae of rows a1, ul, i1, and the anterior part of n, together with a certain
" condensation " of other rows (d1, preoperculo-mandibular, opercular, and other
series) and loss of rows d and b. The pattern of sensory papillae in Lebetus is obviously
more specialized than that in Buenia, but the features present in both denote a
greater affinity between these two gobies than is exhibited between Lebetus and other
genera. In the persistence of cephalic canals and various rows of papillae, Buenia
displays more ostensibly primitive characters than Lebetus. The former in addition
retains the anterior pelvic membrane, is not so small as Lebetus, and does not show
pronounced sexual dimorphism, although this does occur in growth of the first dorsal
fin rays and probably in coloration of this fin. In the male genitalia, the testes are
long and there is no free sperm duct. B. Jeffrey sii may accordingly show greater
resemblance to the common stock from which the two are derived. The Mediterranean
246
P. J. MILLER
species B. reticulatus (C.&V.) (=B. affinis (Kolombatovic)) is more removed from
Lebetus in having a greater number of papillae especially in rows b and d, and a
somewhat higher scale count (more than 33), than B. jeffreysii (Sanzo, 1911, pi. 9,
figs. 9, 10 ; De Buen, 19300). In Manx waters B. jeffreysii has an offshore distribution
FIG. 21. Sensory papillae and orifices of cephalic canals in Buenia jeffreysii, male, 31
mm. standard length. (A) Lateral, (B) dorsal views of head. Abbreviations as in Fig. 7.
Orifices of canals cross-hatched, with greek lettering as in Sanzo (1911).
like Lebetus and has even been taken from the same localities (see also Le Danois,
1910), but differences in precise habitat are perhaps reflected in the different body
form and coloration (Holt & Byrne, 1903, fig. 4).
Some of the characters of Lebetus may be interpreted as adaptations to a mode of life
involving intimate association with dead shells, nodules of calcareous algae, and
stones. The small size of the fish is obviously related to this type of habitat, and the
normal coloration may have cryptic value against a background of Lithothamnion.
Te Winkel (1935) has discussed those anatomical features of the gobiid Mistichthys
TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 247
luzonensis which may be correlated with the extremely small size of this species at
maturity (standard length 10-0-11-0 mm.). Although Lebetus is diminutive, it is
appreciably larger than Mistichthys and the only obvious comparable modification
in this genus is the reduced size of the testis, to which may be related the form of the
urogenital papilla as considered above. A more detailed examination of the viscera
than was attempted in the present study would be needed to investigate this question.
Reduction in number of sensory papillae has already been mentioned, and may be
linked with small body size (Barlow, 19616). The significance of loss of the cephalic
canals is unknown ; these are also lacking in Lesueurigobius, which attains 100 mm.
in L. friesii, but are present in smaller forms such as Odondebuenia, Buenia, etc.
Another feature of probable adaptive importance is the absence of the anterior
pelvic membrane, since this is also missing in Odondebuenia which appears to have a
similar habitat and which in addition exhibits almost complete separation of the pelvic
fins. While the exact affinities of Odondebuenia await investigation, it seems likely that
in view of the resemblances in arrangement of sensory papillae between this genus and
Gobius the condition of the pelvics is derived from an original gobiid disc structure in
response to environmental demands, and is not primitive as in the Eleotrids. The
advantages conferred by the alteration of the pelvic disc in Lebetus and Odondebuenia
are not obvious. The loss of the anterior membrane seems to have occurred inde-
pendantly in the evolution of the two genera as the exploitation of similar ecological
niches proceeded in two areas from different stocks.
It must be stressed that the above phylogenetic conclusions may at present be
regarded as tentative. Not only does the position of Odondebuenia remain doubtful,
but the alignment of Lebetus with Buenia may well be affected by future work on the
lateral line system of exotic gobies and faunistic exploration in little known areas,
which together could disclose closer resemblances and intermediate forms between
these and Lebetus.
SUMMARY
The teleostean genus Lebetus Winther 1877 (Percomorphi-Gobioidea) has been
studied from new Irish Sea material dredged off the south of the Isle of Man, and
from other examples, including types, in the collections of various British and Scandi-
navian museums. A redefinition of the genus includes the disposition of sensory
papillae ; there is a reduction in number of the latter, and cephalic canals and an
anterior membrane to the united pelvic fins are lacking. Two species were previously
recognized : orca Collett 1874 and scorpioides Collett 1874. Among the various
criteria — coloration, meristic characters, body proportions — used in the past to
distinguish these two species, it was found that only in coloration and development of
the dorsal fins could the present Lebetus material be divided into two groups, termed
orca and scorpioides. The former consisted entirely of males, nearly all maturing or
mature, the latter of females and immature males. The conclusion is reached that
there is only one valid species, by page priority Lebetus orca. A detailed account of
the external anatomy and osteology of this species is provided.
In distribution L. orca is restricted to the European Atlantic boreal region, and a
full list of records is provided in an appendix. The species is known chiefly from
248 P. J. MILLER
coarse grounds, and has a wide bathymetric range from about 2 to 375 m. Investiga-
tion of gut contents, using a points system of assessment, shows that L. orca is
exclusively predatory, feeding on small demersal animals which largely comprise
Crustacea and polychaetes. A number of endoparasites are listed. The male repro-
ductive organs are peculiar in the relatively small size of the testis and the long free
sperm duct ; " seminal vesicles " are present. Sexual differences exist in the form
of the urogenital papilla, which in mature males has an unusual terminal circlet of
vascularized papillae. In Manx waters, the breeding season probably commences in
March and may extend to August. 140 to 270 ripening oocytes have been counted in
mature females. Sexual dimorphism is found in coloration, and size of dorsal and
anal fins. Examination of scales indicates that duration of life may be at least two
years, with sexual maturation at one or two years. Maximum total length recorded
is 39-0 mm. Investigation of life history and growth in the Manx specimens was
complicated by the high selectivity of the sampling gear employed.
In a discussion of the systematic position of Lebetus, it is shown that the skeleton is
typically gobiid, and the arrangement of sensory papillae together with other features
suggest that the nearest relative of this genus among the European gobies is Buenia
Jeffrey sii (Giinther). Previous grouping of Lebetus with two Mediterranean genera
lacking an anterior pelvic membrane (Odondebuenia De Buen and Cabotichthys
Whitley) is criticized. The validity of Cabotichthys is doubted, and the suggestion is
made that loss of the anterior membrane in Lebetus and Odondebuenia occurred
independently during their evolution and occupation of a similar habitat in different
areas.
ACKNOWLEDGEMENTS
I wish to thank Messrs J. S. Colman and A. B. Bowers, Marine Biological Station,
Port Erin, and Mr. A. C. Wheeler, British Museum (Natural History), for their
helpful criticism of this paper. My thanks are also due to Mr. D. Eggleston, Dr. R. G.
Hartnoll and Dr. M. C. Miller for collecting specimens of Lebetus while dredging off
the Isle of Man. For arranging the loan of other material, I am most grateful to Mr.
E. K. Barth, Zoologisk Museum, Oslo ; Dr. E. Bertelsen, Carlsbergfondets Dana-
Ekspeditioner, Charlottenlund Slot ; Mr. J. Nielsen, Universitetets Zoologiske
Museum, Copenhagen ; Mr. A. C. Wheeler, British Museum (Natural History) ;
and Mr. J. F. Willgohs, Zoologisk Museum, Bergen. Professor H. Brattstrom,
Biological Station, Espegrend ; Dr. H. O. Bull, Dove Marine Laboratory, Cullercoats;
Mr. B. Christiansen, Zoologisk Avdeling, Tromso Museum ; Dr. R. S. Clarke, Royal
Scottish Museum, Edinburgh ; Mr. S. Mathiasson, Naturhistoriska Museet, Goteborg;
Dr. B. B. Rae, Marine Laboratory, Aberdeen ; Dr. B. Swedmark, Kristinebergs
Zoologiska Station, Fiskebackskil ; and Dr. A. E. J. Went, Fisheries Division of Dept.
of Lands, Dublin, have kindly provided information on distribution or material
present in collections under their care. Messrs J. W. Coles and S. Prudhoe, British
Museum (Natural History), identified nematodes and platyhelminthes encountered in
this study, while Dr. Eve C. Southward, Plymouth Laboratory, and Drs. N. S. Jones
and D. I. Williamson, Port Erin Marine Biological Station, identified polychaetes,
molluscs and arthropods among the gut contents. I am also most grateful to Mr. C. A.
TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 249
Grainge, Zoology Dept., Liverpool University, for preparation of sections, and to
Messrs P. J. Evennett, Zoology Dept., Leeds University, and D. J. Slinn, Port Erin
Marine Biological Station, for photographic assistance. Most of the above investiga-
tion was carried out during the tenure of a D.S.I.R. Research Studentship at the
Marine Biological Station, Port Erin, and has been completed in the Zoology Dept.,
Glasgow University. I am therefore indebted to Mr. J. S. Colman and Professor
C. M. Yonge, F.R.S., for provision of research facilities at Port Erin and Glasgow
respectively.
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TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 253
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APPENDIX
Distribution of Lebetus orca
A list of published and unpublished records known to the author is given below. A complete
sequence of information is as follows : locality, number of specimens, sex, museum number,
standard length + caudal fin length or total length in millimetres (mm.), depth of water in
metres (m.), nature of ground, date of capture, and authority in the case of published records.
When a number of records from the same locality or area are given in the one publication, these
are separated by semicolons and the authority placed at the end. Abbreviations employed are :
c., coarse ground ; cl., clay ; d., damaged; gr., gravel; h., hard ground ; Lith., Lithothamnion ;
m., mud ; midw., midwater ; Mod., Modiolus ; n.d., no date ; s., sand ; sh., shells ; St., stones ;
surf., surface ; t.L, total length ; w., weed.
ICELAND : Off S. and W. coasts, Eystra Horn to BreiSifjordur, " pelagic fry . . . rather rarely "
(Taning, 1940; Saemundsson, 1949).
FAEROES : Off N. coast, and Faero Bank, postlarvae (Taning, 1940).
ROCKALL : On and near Rockall Bank, 1908 (Tailing, 1940).
NORWAY
Hemnefjord (Trondelag) : one vl and pvz) . Two setae are added to the segment in
the deutonymph, one dorsal (d) and one ventral (av). This chaetotaxy is retained
by the adult. The chaetotaxy of trochanter II in the larva and protonymph is similar
to that of trochanter I at those stages but in the deutonymph only one seta (av) is
added to the segment. Four setae occur on trochanter III in the larva and on tro-
chanters III and IV in the protonymph, and five setae on each of these segments in
the deutonymph and adult. The distribution of these setae is more variable than the
setae of trochanters I and II. This is probably due to the asymmetrical development
of the trochanters (and femora) of legs III and IV as the result of the posterior
direction of legs III and IV.
Femora I -IV (Text-figs. 2a-i)
Femora I and II have a completely different chaetotaxy from femora III and IV
as regards the number and the distribution of the setae.
Femur I in the larval and protonymphal stages bears ten setae — four dorsals
(adlt adz, ad3 andpd^, four laterals (allt al2, pl^ and_£/2) and two ventrals (v^ and vz).
The deutonymphal complement of thirteen setae is formed by the addition of one
dorsal (pd2) and two ventrals (v3 and i>4). This number is retained by the adult. There
are no setae situated proximal of the lyriform fissure. Setae pd2 and ad3 assume a
median position on the dorsal surface of the segment.
Femur II in the larva (Text-fig. 2b) has seven setae (ad^ ad2, pdv pd2, vlt al^ and
pli). In the protonymph (Text-fig. 20) a single ventral seta (vz) is added to the seg-
ment whilst in the deutonymph (Text-fig. 2g) three setae appear, one lateral (alz),
one dorsal (ad3) and one ventral (v3] . Setae al2 and v3 are situated proximal to the lyri-
form fissure. The chaetotaxy of the female resembles that of the deutonymph but
in the male the segment becomes swollen and seta v^ and often v2 hypertrophy and
form spur-like structures. The main femoral spur develops from seta vv and the
axillary spur from seta v2. Seta v3 in the male moves to a more distal position and
never lies proximal to the lyriform fissure.
Femur III in the larva and protonymph carries five setae, comprising three dorsals
(adlt ad2 3.ndpd^), one lateral (al) and one ventral (v). Only one seta (pdz) is added to
the segment in the deutonymph. The chaetotaxy of femur IV in the protonymph
resembles that of femur III at that stage except for the absence of the ventral seta
(v) which appears with pdz in the deutonymph. The deutonymphal chaetotaxy of
femora III and IV is retained by the adult.
Genua I-IV (^a-h)
Genu I carries eight setae in the larva and protonymph — four dorsals (adlt ad2,
pdt, pdz), two laterals (a/x and^/j) and two ventrals (av-^ and^Wj). This number is
increased to thirteen in the deutonymph by the appearance of one ventral (avz), two
laterals (alz and plz) and two dorsals (ad3 and pd3). Seta avz migrates medially so
that the three ventral setae form an inverted triangle. Six setae occur on genu II
CHAETOTOXY OF THE LEGS IN THE FREE-LIVING GAMASINA 281
Ki
1.°
-
M
•8.0. "S-°
T3
* ±
~S° -00
0 •>
o
.a
"8°
<
^ lo
-60
-60
^^
o
O
— o
o
o3
1
•JSi
I
CL
-^~
"SP
CL
-«-
CL
-e-
"8°
cr
CL
-»-
CL
-e-
to
CL
-e-
CL
-»-
II3
|| a
d o O
gs.rt
.
>.
OCL
a CL
E
>-
c
o
x
Qo<
ZOOL. 10, 5
I5§§
282
G. OWEN EVANS
°
-DO
o
CL
-©-
•8°-
-50
a
CL
_
-00
°-
a.
•go
M
"8°
A
o"^*
• 2>" «
o 0"°
_A —
o
o
o-S.
Q.
Q.
DO.
CHAETOTOXY OF THE LEGS IN THE FREE-LIVING GAMASINA 283
fel
Q_
-e-
-oO'
o
:>
o
CL
-»-
CL
T3Q •>
O W O
"go
5
Q.
-e-
& *?
O ^
CL
^
^"H"o
O c: £i
__J DCL
1*8
O 4J
Q.
PL,
CO O
h
S
CL.
-e-
•go
.n-tg.
adlt ad2, ad3, pdlf pd2, plv plz> vl and v2
(Text-fig. 66). This type occurs in the genera Sejus, Platyseius and Plesiosejus
(Aceosejidae) and Thinoseius (Eviphididae).
Xll-type (2— f — 2).
Seta vs is added to those occurring in the Xl-type making three ventral setae
(Text-fig. 6c). This chaetotaxy occurs in the families Phytoseiidae and Ameroseiidae,
and in the genera Asca and Halolaelaps (Rhodacaridae) and the genera Lasioseius,
Proctolaelaps, Melichares, Leioseius, Arctoseius, iphidozercon and Zerconopsis
(Aceosejidae).
XHI-type (2-1-2).
This type is characterized by having four ventral setae (Text-fig. 6d). It is the
most widely distributed chaetotactic pattern in the suborder and has been observed
in the families Parasitidae, Veigaiaidae, Arctacaridae, Laelaptidae, Zerconidae,
Epicriidae, Macrochelidae and Pachylaelaptidae, and in the genera Eviphis, Alliphis,
Pelethiphis and Scardbaspis (Eviphididae), and Rhodacarus, Rhodacarettus, Eury-
parasitus, Gamasellus, Ologamasus and Gamasiphis (Rhodacaridae).
CHAETOTbXY OF THE LEGS IN THE FREE-LIVING GAMASINA 287
al,
al,
ad,
av,
av2
°H*°
ad2
a
pi.
a I,
Ol:
o
) ad,
pdi
o
V2
c
o
ad2
> ad3
• o3
Vi
pd2
c
pl:
al,
ad,
o
pd,
o .
ad2
o
(
V2
v,
pd2
o
ad3
V3 (
pi.
al,
o Pd
ad, °
9
ad,
a
ad3
o
op!2
FIGS. £>Or-d. Diagrammatic representation of the chaetotaxy of femur I in the free-living
Gamasina, a, X-type (Microsejidae). b, Xl-type (Aceosejidae). c, Xll-type (Amero-
seiidae). d, Xlll-type (Eviphididae) .
Abbreviations as in Fig. i.
Genu I
I have observed five types of chaetotactic patterns of germ I ; the number of
setae ranging from ten to thirteen.
288
G. OWEN EVANS
X-type (2-f , f-2).
The following setae are present : adv ad2, pdlt pdz, avlt pvlt alv al2, pllt plz (Text-
fig. 7«). This type is characteristic of members of the families Phytoseiidae and
Microsejidae.
ad,
ad,
all <
o
) • Pd,<
>pl|
o pd.
^\
av, •(}
PV.
al| <
) a;' ,° <
pi,
\j
ad2
f^
o
a!2<
> ad2
>p!2 al2<
pd,
§f^i_
UI2
pd2
ad,
°av, pd,
ad3
a al'(
• pv,
av2 r
pi, b
0
ad2 0
>PI2
ad,
ad3
ao' Pdl
O ,
o
ah <
> • ^o' <
>pl, Q all <
av.
ipll
av, •
O pv,
pVi
Of
ad2 • o
2 pd2
a!2 i
' o l °d '
^P^ a bi
o pd3
>pk
ad, pdl
ads O
FIGS. ya-«. Diagrammatic representations of the chaetotaxy of genu I in the free-living
Gamasina. a, X-type (Phytoseiidae) . b, Xlb-type (Macrochelidae) . c, XIa-type (Evi-
phididae). d, XH-type (Aceosejidae). e, XHI-type (Parasitidae).
Abbreviations as in Fig. i.
XIa-type (i-|, f-2).
This chaetotactic pattern differs from all others on genu I in having a single
CHAETOTOXY OF THE LEGS IN THE FREE-LIVING GAMASINA 289
antero-lateral (al-^ seta. The setal complement is alv adv adz, adz, pdv pd2, plv plz,
avlt avz and pv^ (Text-fig. 7^) . I have only observed this pattern in the Eviphididae.
Xlb-type (2-f, f-2).
This differs from the X-type in having an additional dorsal seta, adz (Text-fig. 7&).
It is characteristic of the Macrochelidae.
XH-type (2-|, |-2).
This chaetotactic pattern is similar to Xlb except for the presence of an additional
ventral seta (avz). The three ventral setae form an inverted triangle owing to the
migration of seta avz to a median position on the segment (Text-fig. 7^). It is found
in the Ameroseiidae and in the genera Arctoseius, Iphidozercon and Zerconopsis
(Aceosejidae), Asca, Digamasellus and Halolaelaps (Rhodacaridae) and Pachyseius
(Pachylaelaptidae) .
Xin-type (2-f, f-2).
The XUI-type is characterized by having six dorsal (ad-^-ad^ and pdf-pda), in
addition to two antero-laterals, two postero-laterals and three ventrals (Text-fig. 70).
I have observed this genual pattern in the families Parasitidae, Veigaiaidae, Arcta-
caridae, Laelaptidae, Zerconidae and Epicriidae, and in the genera Lasioseius
Proctolaelaps, Melichares, Leioseius, Sejus, Platyseius and Plesiosejus (Aceosejidae),
Pachylaelaps and Olopachys (Pachylaelaptidae) and Rhodacarus, Rhodacarellus,
Euryparasitus, Gamasellus, Ologamasus and Gamasiphis (Rhodacaridae).
Tibia I
The X-XIII-types of chaetotaxy observed on genu I also occur on tibia I as well
as a XlV-type characterized by having four ventral setae.
X-type (2-f, f-2).
This type (Text-fig. 8a) is found in the families Phytoseiidae and Microsejidae.
Xl-type (i-f , f-2).
Only the XIa-type of chaetotaxy of genu I occurs on the tibia and as in the case of
the genu it appears to be restricted to members of the family Eviphididae (Text-fig.
86).
XH-type (2-|, f-2).
This type (Text-fig. 8c) occurs in the families Macrochelidae, Pachylaelaptidae
and Ameroseiidae, and in the genera Arctoseius, Iphidozercon, Zerconopsis and
Plesiosejus (Aceosejidae), and Digamasellus and Halolaelaps (Rhodacaridae).
XIH-type (2-f, f-2).
I have observed this chaetotactic pattern (Text-fig. 8d) in the free-living Laelaptidae
and in the genera Lasioseius, Proctolaelaps, Melichares, Leioseius, Sejus, Platyseius
and Plesiosejus (Aceosejidae), and Asca (Rhodacaridae).
2QO
G. OWEN EVANS
a!2o
ad,
o
pa.
Opli
°A
ad2
a
al.
a!2o
o'
av, pv.
opli
ad
t— %
ad,
al,
pd,
o
O
ad2 o
pi,
ad n
a!
ad i
0 pd,
av, 0
av2
o
ad2
ads
0
pd2
PV.
q
ad3 o
p'd
FIGS. 8a-e. Diagrammatic representations of the chaetotaxy of tibia I in the free-living
Gamasina. a, X-type (Phytoseiidae) . b, Xl-type (Eviphididae). c, XH-type (Aceoseji-
dae). d, XHI-type (Aceosejidae). e, XlV-type (Parasitidae) .
Abbreviations as in Fig. i.
XlV-type (2-|, |-2).
The four ventral setae arranged in the form of a rectangle readily distinguishes this
pattern from the others found on tibia I (Text-fig. Se). It occurs in the families Para-
sitidae, Veigaiaidae, Arctacaridae, Zerconidae, Epicriidae and in the genera Rhoda-
carus, Rhodacarellus , Cyrtolaelaps, Euryparasitus , Gamasellus, Ologamasus and
Gamasiphis (Rhodacaridae) .
CHAETOTOXY OF THE LEGS IN THE FREE-LIVING GAMASINA 291
O.
-e-
a.
-e-
-oO
a
(J
(3P
O ™
•>-> o
^> c3
^Tab
If 8,
s*£>
s~~
4) f-H JH
^ £
rt «J > «
O ^H 'd *t3 W "s
2Q2 G. OWEN EVANS
Femur II
The following four types of chaetotactic pattern have been observed on this
segment :
IX-type (2-f-i).
This type has the following chaetotaxy ; alv, alz, adv adz, pdlt pd2, plv av{ and pv^
(Text-fig. go). It occurs in the Microsejidae.
Xa-type (2-|-i).
The chaetotaxy comprises adv adz, pdlt pdz, alv alz, pllt vlt v2 and vz. This type
has been found in the family Ameroseiidae.
Xb-type (2-f-i).
The chaetotaxy differs from the Xa-type in the presence of five dorsal setae (ads
additional) but only two ventral setae. It is characteristic of the Phyto-
seiidae, the genera Sejus, Platyseius and Plesiosejus (Aceosejidae), and also occurs
in the genus Thinoseius (Eviphididae). In the latter, the ventral setae are widely
separated so that one occurs in the proximal and the other in the distal half of the
segment (Text-fig, gc). In the Phytoseiidae, however, both ventral setae are usually
situated in the proximal half of the segment (Text-fig, gb).
Xl-type (2-|-i).
This type has five dorsal setae and three ventrals in addition to the two antero-
laterals and one postero-lateral (Text-fig, gd). It is the most widely distributed
chaetotactic pattern of femur II in the Gamasina and has been observed in the families
Parasitidae, Veigaiaidae, Arctacaridae, Rhodacaridae, Zerconidae, Epicriidae,
Laelaptidae, Macrochelidae, Pachylaelaptidae, Aceosejidae (with the exception of
Sejus, Platyseius and Plesiosejus) and Eviphididae (excluding Thinoseius).
Seta al2 is always situated proximal to the lyriform fissure in all four types of
chaetotaxy.
Genu II
Genu II carries seven, nine, ten or eleven setae.
VH-type (2-§, g-i).
In this type there are no ventral setae (Text-fig. 100) ; the chaetotaxy comprising
setae allt alz, adlt ad2, pdlt pd2 and plv It occurs in the majority of the Phytoseiidae.
IX-type (2-f , f-i).
I have, to date, only observed this pattern in Amblyseius graminis Chant (Phyto-
seiidae). The chaetotaxy shows an addition of two ventral setae (av-^ and pvj to
the VH-type (Text-fig. io&).
Xa-type (2-f, §-2).
This differs from the IX-type in having two postero-lateral setae (Text-fig. 100).
This pattern is characteristic of the Microsejidae.
CHAETOTOXY OF THE LEGS IN THE FREE-LIVING GAMASINA 293
a!2 (
a!2 (
ad,
> ad,
pd,
O
o<
pd2
ipl
ali I
ad,
O
pd,
o
ad,
O
pd,
0
N av,
, i
c
i
av,
pv* U '
•
o
ad2
O
nM
> o
at<
) _
pd2 (
ad2
pd2
O
ad3
a
ad,
0
pd,
0
pv,
2
Q
'
fi pv,
pd2
Qi
ad3
FIGS. loa-f. Diagrammatic representations of the chaetotaxy of genu II in the free-
h'ving Gamasina. a, VH-type (Phytoseiidae). b, IX-type (Amblyseius graminis Chant :
Phytoseiidae) . c, Xc-type (Sejus : Aceosejidae). d, Xb-type (Aceosejidae). e, Xa-type
(Microsejidae). /, XI -type (Parasitidae) .
Abbreviations as in Fig. i.
Xb-type (2-f, 5-i).
This type (Text-fig. iod) which occurs in the genera Arctoseius, iphidozercon and
Zerconopsis (Aceosejidae) is characterized by having two antero-laterals (al^ and al2),
five dorsals (ad^3 and pd^), two ventrals (av^ and PV-^) and one postero-lateral
Xc-type (2-f, §-2).
The Xc-type has two postero-laterals but only one ventral seta (Text-fig. loc).
It occurs in the genera Sejus, Platyseius and Plesiosejus (Aceosejidae).
Xd-type (i-f , |-2).
This pattern differs from all others on genu II in having only one antero-lateral
seta. I have observed this chaetotaxy in the genus Thinoseius (Eviphididae) only.
294 G- OWEN EVANS
XI-type (2-1 f-2).
The XI-type (Text-fig. io/) is by far the most widely distributed chaetotactic
pattern of this segment and occurs in the families Parasitidae, Veigaiaidae, Arcta-
caridae, Rhodacaridae, Zerconidae, Epicriidae, Macrochelidae, Pachylaelaptidae,
Laelaptidae, Ameroseiidae and Eviphididae (excluding Thinoseius) and in the genera
Lasioseius, Leioseius, Melichares and Proctolaelaps (Aceosejidae).
Tibia II
In the species I have examined, tibia II bears seven, nine or ten setae.
VH-type (i-f , i-i).
This type is characteristic of the Phytoseiidae and comprises setae ah, adv adz, pdlt
plv av! andpvi (Text-fig,
IXa-type (i-f , f-2).
This type, observed only in Thinoseius (Eviphididae), has one antero-lateral seta
K).
IXb-type (2-1 f-i).
The chaetotaxy comprises allt alz, adv adz, pdlt pd2, plv av^ andpv2 (Text-fig. n&).
This type occurs in the genera Arctoseius, iphidozercon and Zerconopsis (Aceosejidae).
IXc-type (2-f , 1-2).
The chaetotaxy comprises setae alv alz> adv adz, pdv pllt pl2, avlf pv± (Text-fig, nc).
I have observed this type in the genera Platyseius and Plesiosejus (Aceosejidae).
IXd-type (2-1, |-2).
This differs from the IXc-type in the setation of the dorsal surface of the segment ;
the three setae being adlt pd1 and pd2 (Text-fig, n^). It occurs only in the Micro-
sejidae.
X-type (2-f, f-2).
The following setae are present : allt alz, ad^ adz, pdv pd2, plv pl2, av^ and pvt
(Text-fig. lie). This chaetotactic pattern is present in the families Parasitidae,
Veigaiaidae, Arctacaridae, Rhodacaridae, Zerconidae, Epicriidae, Laelaptidae,
Ameroseiidae, Macrochelidae, Pachylaelaptidae, Eviphididae (excluding Thinoseius)
and in the genera Lasioseius, Leioseius, Melichares, Proctolaelaps and Sejus (Aceoseji-
dae).
Tarsi II-IV
The chaetotaxy of tarsi II-IV is extremely constant throughout the free-living
Gamasina ; the details given for Pergamasus (p. 278) being applicable throughout.
There is some variation in the relative position of the setae due, in some instances,
to the elongation of the segment and in the form of setae ad± and pd^.
CHAETOTOXY OF THE LEGS IN THE FREE-LIVING GAMASINA 295
al,
al,
al2*
'ad,
O
pd,
O
av,
1
A<
O
ad.
av, pv,
»
Pd
adi
pd,
O
> av,
(
PV,
O
pd2
O
\
ad 2
FIGS, iia-e. Diagrammatic representations of the chaetotaxy of tibia II in the free-
living Gamasina. a, VH-type (Phytoseiidae) . b, IXb-type (Zerconopsis : Aceosejidae).
c, IXc-type (Platyseius : Aceosejidae). d, IXd-type (Microsejidae). e, X-type (Laelap-
tidae).
Abbreviations as in Fig. i.
Femur III
The number of setae on femur III is markedly constant. With the exception of
Thinoseius, all the species I have examined have six setae (Text-fig, ge). Owing to the
difference in shape of this segment in certain species, the distribution of the setae is
subject to some variation. In Thinoseius, seven setae are present ; the additional
seta apparently being developed ventrally (Text-fig, o,/).
Genu III
This segment bears six to ten setae.
296 G. OWEN EVANS
Vl-type (i-g, §-i).
The following setae are present : alv adlt adz, pdv pdz and plt (Text-fig. I2«).
There are no ventral setae. I have seen this type only in the genus Phytoseius
(Phytoseiidae).
VH-type (i-», g-i).
The chaetotaxy comprises setae allt adlt ad2> pdlt pdz, pl± and av^ (Text-fig. 126).
It occurs in members of the family Macrochelidae and in the genera Arctoseius,
Iphidozercon and Zerconopsis (Aceosejidae), Pachylaelaps and Olopachys (Pachylae-
laptidae), Thinoseius (Eviphididae) and Iphiseius and Typhlodromus (Phy toseiidae) .
Villa-type (i-J, f-i).
The chaetotaxy differs from the VH-type in the addition of pv-L (Text-fig. I2c).
This chaetotactic pattern occurs in the Eviphididae (excluding Thinoseius).
VHIb-type (2-1 8-i).
This chaetotactic pattern has been found in the genera A sea and Saprolaelaps
(Rhodacaridae), Sejus, Platyseius and Plesiosejus (Aceosejidae), Pachyseius (Pachy-
laelaptidae) and Epicriopsis (Ameroseiidae). The eight setae comprise ah, al2, adv
adz> pd±, pdz, pl± and avt (Text-fig, xzd}.
IXa-type (2-}, f-i).
Setae o/lf alz, adv ad2, pdv pdz, pllf av± and pvl are present in this type (Text-fig.
120). This pattern occurs in the families Parasitidae, Veigaiaidae, and in the genera
Rhodacarus, Rhodacarellus, Cyrtolaelaps, Euryparasitus, Gamasellus, Hydrogamasus,
Ologamasus and Gamasiphis (Rhodacaridae), Lasioseius, Leioseius, Melichares and
Proctolaelaps (Aceosejidae), Hypoaspis, Coleolaelaps and Ololaelaps (Laelaptidae)
and Neocypholaelaps (Ameroseiidae).
IXb-type (2-f, §-2).
I have observed this type only in the Microsejidae (Text-fig. I2/).
X-type (2-|, ?-2).
The ten setae comprise allt al2, ad^, ad2, pdlf pdz, pllt plz, av^ and^Vj (Text-fig. ~L2g}.
It occurs in the families Arctacaridae, Zerconidae and Epicriidae, and in the genera
Ameroseius and Kleemannia (Ameroseiidae).
Tibia III
The following three types of chaetotactic pattern have been observed on this
segment :
VH-type (i-l, }-i).
This type occurs in the families Phytoseiidae, Eviphididae, Macrochelidae and
Pachylaelaptidae, and in the genera Arctoseius, Iphidozercon and Zerconopsis
(Aceosejidae). The chaetotaxy comprises a/1} adv pdv pd2, pllt av± and pv± (Text-fig.
CHAETOTOXY OF THE LEGS IN THE FREE-LIVING GAMASINA 297
ad, pd,
ad, pd,
0 O
O O
a!, <
)
c
>pl,
al, <
av,
I
o o
O 0
ad2 pd2
ad2 pd2
/a
b
ad, pd,
o o
• •
aU
ad, pd,
> • °
ad (
av, pv,
)
ipl.
av, (
>pl|
oQ°d' P°d2
c
d
ad,
av, py,
i
<
0 pd,
• _ o
H
al,(
ad, pd,
) O O (
A
»p|, al, <
o0 I
> ad, pd, <
all i
) av, pv,
w
^i
avi
a!2<
o
ad2 o
> Pd2
a!2<
> o o
ad2 pd2
1C 1 ^2 '
>0, 9C
aa2 pa2
pi
FIGS. i2a-g. Diagrammatic representations of the chaetotaxy of genu III in the free-
living Gamasina. a, Vl-type (Phytoseius : Phytoseiidae) . b, VH-type (Macrochelidae) .
c, Villa-type (Eviphididae) . d, VHIb-type (Asca : Rhodacaridae) . e, IXa-type
(Parasitidae). /, IXb-type (Microsejidae). g, X-type (Zerconidae) .
Abbreviations as in Fig. i.
2Q8
G. OWEN EVANS
ad,
pd,
o
O
al,<
) av,
£
pv, (
0
pd2
ad,
0
pd,
O
al, <
.
> av,
PV, <
O
a)2<
>
pd,
ad.
) 0
pd,
0
* <
av,
pv,
)
o (
pd2
FIGS. i3«-c. Diagrammatic representations of the chaetotaxy of tibia III in the free-
living Gamasina. a, VH-type (Phytoseiidae) . b, VHI-type (Parasitidae) . c, IX-type
(Microsejidae).
Abbreviations as in Fig. i.
VHI-type (2-\, f-i).
There is an addition of one antero-lateral seta to the VH-type ; the seta being
tf/2 (Text-fig. 136). This pattern is present in the families Parasitidae, Veigaiaidae,
Rhodacaridae, Laelaptidae, Ameroseiidae (excluding Kleemannia) and in the genera
Lasioseius, Leioseius, Melichares and Proctolaelaps (Aceosejidae).
IX-type (2-1, f-2).
Seta_/>/2 is additional to the VHI-type (see Text-fig. I3c). It occurs in the families
Arctacaridae, Zerconidae, Epicriidae and Microsejidae and in the genus Kleemannia
(Ameroseiidae).
Femur IV
The presence of six setae on this segment is extremely constant in the free-living
Gamasina ; the only exception being members of the Microsejidae. In the majority
of the species I have examined the chaetotaxy comprises alv adv ad2, pdlt pi and vi
(Text-fig, gh) but there are variants of this pattern as, for example, in the aceosejid
Plesiosejus (Text-fig. gg) which has four dorsals but no postero-lateral setae. The
microsejids have seven setae on this segment (alv ad^, ad2, pdv pd2, av1 and^vj.
Genu IV
Five types of chaetotaxy of genu IV have been observed.
Vl-type (i-f, §-o).
The following setae are present : alit adlt ad2, pdv pdz and av± (Text-fig. 140;).
This pattern is found in certain species of the genus Macrocheles s.lat. (Macrochelidae)
and has been used as a taxonomic character by Evans & Hyatt (1962).
CHAETOTOXY OF THE LEGS IN THE FREE-LIVING GAMASINA 299
VII-type (i-f, g-i).
This shows an addition of seta pi to the Vl-type (Text-fig. 146). It is found in the
families Pachylaelaptidae, Eviphididae and Phytoseiidae, in the genera Arctoseim,
iphidozercon and Zerconopsis (Aceosejidae), Digamasellus (Rhodacaridae) and certain
species of the Macrochelidae.
ad, pd,
o o
al, o av,
al,
ah
ad,
o
pd,
0
) •
av,
) O
p32 <
J
od2
o
al
,
al, <
odi pd,
> o o
•
a!2<
c
> o Q
ad2 pd2
ali
a!2
ad,
av,
,
ad