J. Parasitol., 91(1), 2005, pp. 131–134
? American Society of Parasitologists 2005
HAEMOPROTEIDS OF THE AVIAN FAMILY DICRURIDAE (THE DRONGOS)
Amy F. Savage and Ellis C. Greiner*
Department of Pathobiology, College of Veterinary Medicine, University of Florida, Box 110880, Gainesville, Florida 32611. e-mail:
the presence of hematozoa and 1 species of Haemoproteus, i.e., Haemoproteus dicruri (De Mello, 1935). One species of drongo
occurs in Madagascar and has not been examined previously. Blood smears collected from wild-caught crested drongos, Dicrurus
forficatus, in Madagascar were examined using a compound microscope for the presence of hematozoa. A new species, Hae-
moproteus khani, is described in this study. This new species has circumnuclear gametocytes, in contrast to the halteridial H.
dicruri. In addition, H. dicruri is reported for the first time from the crested drongo and is redescribed. This is the first report of
hematozoa in drongos of Madagascar.
Dicrurids are a widespread avian family in Africa and Asia. Earlier surveys of this family in these areas have reported
The Dicruridae includes 20 species of birds, occurring in Af-
rica, India, and Australia (Langrand, 1990). The crested drongo
(Dicrurus forficatus) is the only member of the family on Mad-
agascar; it occurs there and on Anjouan in the Comoro Islands.
Dicrurids are common and can be found in a variety of habitats,
from forests to sparsely wooded terrain and plantations (Morris
and Hawkins, 1998). To date, 3 studies have examined birds
from Madagascar for hematozoa (Bennett and Blancou, 1974;
Greiner et al., 1996; Raharimanga et al., 2002).
As yet, only a single species of Haemoproteus has been re-
ported in the Dicruridae. Haemoproteus dicruri was first de-
scribed by De Mello (1935) from Dicrurus macrocercus and
later redescribed by Peirce (1984) from Dicrurus adsimilis.
Several authors have discussed host specificity in this genus
(Atkinson, 1986; Bennett and Peirce, 1988). For example, At-
kinson (1986) showed that Haemoproteus meleagridis, although
primarily observed in turkeys, was transmissible to other mem-
bers of the Phasianidae. The new Haemoproteus species de-
scribed in this study is distinct from H. dicruri and, to our
knowledge, is specific to the Dicruridae.
MATERIALS AND METHODS
All birds were mist-netted at field sites in Madagascar. Blood smears
were made either from toe clippings or from heart punctures. The slides
were air-dried and fixed in absolute methanol by the collecting orni-
thologists. Slides were either stained before shipment or shipped to the
University of Florida, where they were stained with Giemsa on arrival.
Slides were examined at ?100, ?160, and ?1,000 on a Zeiss compound
microscope for the presence of hematozoa. Blood parasites were drawn
with the aid of a drawing tube on a Nikon compound microscope. All
measurements, excepting parasite length, of erythrocytes and parasites
were performed as described in Bennett and Campbell (1972). Nuclear
displacement ratio (NDR) refers to the degree to which the host cell
nucleus is laterally displaced by the parasite. Nuclear displacement was
calculated as described in Savage and Greiner (2004). Parasite length
was determined by measuring a line drawn to bisect the gametocyte
along its longitudinal axis. Area was calculated using a drawing tube
and grid, as described in Forrester et al. (1977).
Haemoproteus khani n. sp.
(Figs. 1–6; Table I)
Immature gametocyte: Developmental stages halteridial, de-
veloping laterally in mature erythrocyte, with ends wrapping
Received 26 April 2004; revised 17 June 2004; accepted 18 June
* To whom correspondence should be addressed.
around erythrocyte nucleus (Figs. 1, 2). First circumnuclear
contact between parasite ends occurs adjacent to host cell nu-
cleus (Fig. 3), then parasite grows outward, filling remaining
host cell cytoplasm at maturity. Host–parasite complex at this
stage is 10 ?m2larger than the fully mature complex, but par-
asite has same area, indicating host cell atrophy as parasite ma-
Macrogametocyte (n ? 15): Circumnuclear at maturity (Fig.
4), lightly staining blue with Giemsa. Nucleus commonly in-
distinct, light pink when visible. Margins smooth or slightly
irregular, but not amoeboid. Fully developed host cell–parasite
complex area ? 130.5 ?m2, 8% larger than uninfected eryth-
rocytes. Host cell nucleus displaced slightly, NDR ? 0.81. Av-
erage parasite area ? 109.3 ?m2, host cell nucleus reduced from
22.4 to 18.0 ?m2. Parasite length ? 28.8 ?m. Infected cells
slightly increased in length. When visible, parasite nucleus ?
10.4 ?m2(9.5% of parasite). Refractile granules located ran-
domly through parasite, sometimes clumped. Pigment granules
vary in size, generally moderately large and ovoid, occasionally
smaller. Pigment white or yellow. Average number ? 13. No
volutin observed. Parasite develops in mature erythrocytes.
Microgametocyte (n ? 11): Same morphological character-
istics as described above, with gender staining associated dif-
ferences (Figs. 5, 6). Parasite stains pale pink or clear, com-
monly noticed only because of pigment granules. Parasite nu-
cleus pink when observed, average area ? 24.1 ?m2, 22.1% of
Type host: Dicrurus forficatus, Linnaeus, 1766.
Basis of description: Parasites are described from a blood
smear taken from an adult D. forficatus. Hapantotype: blood
smear SG-604 collected by Steven M. Goodman on 10 Decem-
ber 1995 in Andohahela, Madagascar, at 120 m. Accession
G463732 to the International Reference Centre for Avian He-
matozoa, Queensland Museum, South Brisbane, Queensland,
Australia. Parahapantotype: blood smear SG-605 collected by
Steven M. Goodman on 10 December 1995 in Andohahela,
Madagascar, at 120 m. Accession to IPM-11 to Institut de Pas-
teur de Madagascar.
Distribution: It is presumed that this parasite will be found
throughout the range of this species of drongo on Madagascar
and perhaps the Comoro Islands.
Etymology: The parasite is named in recognition of the sig-