Filarioid nematodes (Filarioidea: Onchocercidae) of Corvus brachyrhynchos brachyrhynchos Brehm in...

FILARIOID NEMATODES (FILARIOIDEA: ONCHOCERCIDAE) OF CORVUS BRACHYRHYNCHOS BRACHYRHYNCHOS BREHM IN SOUTHERN ONTARIO, CANADA AND A CONSIDERATION OF THE EPIZOOTIOLOGY OF AVIAN FILARIASIS

Cheryl M. BARTLETT and Roy C. ANDERSON

Department of Zoology, College of Biological Sciences, University of Guelph, Guelph, Ontario, Canada

Summary

Chandlerella chitwoodae Anderson, 1961 (prevalence 65%), Splendidofilaria caperata Hibler, 1964 (21%), Eufilaria longicaudata Hibler, 1964 (15.5%), C. quiscali (von Linstow. 1904) Robin- son, 1971 (1.9%), Cardiofilaria pavlovskyi Strom, 1937 (0.4%) and Splendido]ilaria wehri (0.3%) were found in crows (Corvus brachyrhynchos brachyrhynchos Brehm) wintering in southern Ontario, Canada. S. caperata, E. longicaudata, Cardiofilaria pavlovskyi and S. wehri are reported for the first time from this host.

Infection with Chandlerella chitwoodae and E. longicaudata produced microfilaraemias and these two common species apparently are maintained in the crow population. Infection with S. caperata was amicrofilaraemic (occult) and this common species cannot, therefore, be maintained in the crow population. C. quiscali, Car- diofilaria pavlovskyi and S. wehri occurred infrequently and must be common parasites of other woodland bird species. The presence of both common and sporadic infections in crows sug- gests that some avian filarioids parasitize numerous birds sharing a particular habitat. However, some host species are probably more important than others in maintaining infections within an avian community.

The hosts of each species in ChandlereUa, Splendidofilaria, Cardiofilaria and Eufilaria are listed. Chandlerella flexivaginalis (Jones, 1961) Sultana, 1962 and C. hispanica L6pez- Caballero, 1974 are placed in synonymy with C. chitwoodae. Splendidofilarioides Freitas & Ibfifiez, 1968 is made a synonym of Splen- didofilaria and the type species becomes Splen- Systematic Parasitology 2, 77-102 (1980). 0165-5752/80/0021-0077 $05.20 @ Dr. W. Junk b.v. Publishers, The Hague. Printed in Northern Ireland

did@laria pachacuteci (Freitas & Ibfifiez, 1968) n. comb. S. passerina Koch & Huizinga, 197l is synonymized with S. algonquinensis (Anderson, 1955) Anderson, 1961. Pseudaproctella andersoni var. major Chabaud, Brygoo & Richard, 1964 and Cardiolilaria chabaudi Dissanaike & Fernando, 1965 are placed in synonymy with C. major Dissanaike & Fernando, 1965. Eufilaria cypseli (Annett, Dutton & Elliot, 1901) Nelson, 1966 is transferred to Lemdana as L. cypseli (Annett, Dutton & Elliot, 1901) n. comb. E. buckleyi (Desmukh, 1968) n. comb. is designated for Neofilaria buckleyi Desmukh, 1968 (= Eul'ilaria utae Anderson & Bain, 1976).

Introduction

Fifteen genera of avian filarioids are recognized by Anderson & Bain (1976). These authors point out that some species, particularly those in the larger genera, are possibly synonyms and that many species, in addition to Cardiofilaria pavlovskyi Strom, 1937 and Aproctella stoddardi Cram, 1931, might have wide host and geographical distributions. Unfortunately, many species lack modern descriptions and this has limited our concepts of the numbers and distribution of avian filarioids. The epizootiology of avian filariasis is also poorly understood. Few studies have examined sufficient numbers of a particular bird species to determine whether the filarioids found occur commonly or sporadically in that population. The type host in the taxonomic sense might not be the main host in the epizootiological sense.

Crows (Corvus brachyrhynchos brachyrhynchos

77

Filarioid nematodes of Corvus brachyrhynchos brachyrhynchos Brehm

Brehm) congregate in large winter flocks in southern Ontario, Canada and can be collected easily in large numbers. This offered an oppor- tunity for a detailed study of the filarioid parasites of this species. This paper redescribes the six species of filarioids found and considers their epizootiological relationships. Two species apparently can be maintained in the crow population. The other four species are probably acquired from other infected host species and not maintained in the crow population. Three of these species occurred infrequently and one species commonly; this last always produced occult infections. These observations suggest that avian filarioids tend to occur more within bird communities than within particular host species.

Methods and materials

Crows were attracted to offal placed in a field in Essex County, southern Ontario, Canada, during the winters 1977-78 and 1978-79. They were then either shot by hunters concealed in nearby blinds or live-trapped with a rocket net-trap. Live crows were subsequently killed by decapita- tion. Carcasses of crows were examined with a stereoscopic microscope for adult filarioid nematodes. Adult worms recovered were fixed in glycerin-alcohol and cleared for examination in glycerin.

Microfilariae in the blood of the lungs were associated with adult worms by comparing their morphology with that of microfilariae found in the vagina of living female worms recovered from the tissues of the crows.

Microfilariae were vitally stained following a modification of the method of Schillhorn van Veen & Blotkamp (1972). A drop of 1% al- coholic solution of brilliant cresyl blue was placed on a glass slide and spread in a manner similar to that of a thin blood smear. One or more drops of physiological saline and a drop of lung blood were placed in the centre of the slide, mixed and covered with a vaseline-ringed cover slip. The positions of certain features (i.e. nerve ring, excretory vesicle, excretory cell, inner body, rectal cells and anal vesicle) were determined three to six hours later.

The technical reference used for the taxonomy of bird species was Peters (1931-62).

Results, redescriptions and lists of species

The prevalence of filarioid nematodes in crows found in this two-year study in Southern Ontario is shown in Table I. The six species found, namely, Cardiofilaria pavlovskyi, Chandlerella chitwoodae, C. quiscali, Splendidofilaria caperata, S. wehri and Eufilaria longicaudata are rede- scribed below, and the valid species in the four genera are listed. The host distribution reported is tabulated (see Tables II to V).

1. Cardiofilaria pavlovskyi Strom, 1937 (Figs. 1-8, Table I)

Synonyms: Ornithofilaria inornata Anderson, 1956; Carinema ardae Mawson, 1957; Pseuda- proctella inornata (Anderson, 1956) Anderson & Chabaud, 1959; Pseudaproctella ardae (Mawson, 1957) Anderson & Chabaud, 1959; Cardiofilaria inornata (Anderson, 1956) Sonin, 1961; C. ardae (Mawson, 1957) Sonin, 1961.

General: Filarioidea, Onchocercidae, Splen- didofilariinae, Cardiofilaria Strom, 1937. Slender nematodes with bluntly rounded extremities. Cuticle thin and smooth. Cephalic extremity with

Table I

Prevalence* of filarioid infections in crows (Corvus brachyrhynchos brachyrhynchos Brehm) collected in Essex County, southern Ontario, during the winters of 1977-78 and 1978-79.

No. crows No. crows Prevalence Species examined infected %

Cardiofilaria pavlovskyi 269 1 0.4 Chandlerella chitwoodae 269 174 65.0 Chandlerella quiscali 269 5 1.9 Splendidofilaria caperata 341 72 21.0 Splendidoj'ilaria wehri 341 1 0.3 Eufilaria longicaudata 110 17 15.5

* Prevalence of C. chitwoodae and E. longicaudata based on presence of microfilariae in lung blood; data do not include non-pa ten t or single sex infections. Prevalence of (7. pavlovskyi, C. quiscali, S. caperata and S. wehri based on presence of adult nematodes ; data do include non-pa ten t or single sex infections.

78

Table II

Host distribution by avian family of species of Cardiofilaria Strom, 1937"

E

Avian Order "~ ~ "~ ~ ~ -~ -- g E and Family ~ ~ ~ ~-~ E ~= E .~

ARDEIFORMAES Ardeidae

ACCIPITRIFORMES Acciptridae Falconidae

GALLIFORMES Phasianidae

CHARADRIIFORMES Charadriidae Scolopacidae

PSITTACIFORMES Psinacidae

CUCULIFORMES Cuculidae

STRIGIFORMES Strigidae

APODIFORMES Apodidae x

PASSERIFORMES Formicariidae Oriolidae Corvidae Campephagidae Muscicapidae Motacillidae Laniidae Nectariniidae Parulidae

X

×

×

X

)<

)<

ixl

× ×

* All species reported from the body cavity, pavlovskyi also reported from the heart cavity, and maior from the subcutaneous tissues.

four pairs of submedian papillae and lateral amphids. Oral opening small and circular. Buccal cavity surrounded by delicate cuticular ring. Oesophagus short, well developed and clearly demarcated from broad intestine, not externally divided into muscular and glandular parts. Phas- mids terminal.

Male (25 specimens): Length 7.6 (6.5-8.8) ram. Maximum width 135 (105-184) ~m near middle of body. Nerve ring 165 (140-194) t~m from anterior extremity. Oesophagus 220 (176-250) txm long and 28 (15-40) ixm wide. Caudal extremity coiled. Tail 73 (56-94) txm in length.

C.M. Bartlett & R.C. Anderson

Five to seven minute, circumanal papillae present. Spicules subequal and dissimilar, right 65 (54-76) txm long with rounded distal end, left 76 (64-84) t~m long with trough-shaped distal end.

Female (25 specimens): Length 14.1 (10.8-19.1) mm. Maximum width 292 (235-370) txm near middle of body. Nerve ring 202 (155-260) ~m from anterior extremity. Oesophagus 528 (410- 675) t~m long and 30 (20-45) Izm wide. Vulva 1281 (775-1550) izm from anterior end of body, slightly salient. Vagina 818 (625-950) t~m long, extending short distance posteriorly, then bending anteriorly to pass vulvar region before turning posteriorly and joining uteri. Didelphic and opisthodelphic. Tail 183 (125-260) txm in length. Anus difficult to discern. Posterior extremity generally with two large lateral swellings.

Microfilaria (20 specimens from the vagina): Length 350 (270-400) txm. Width of anterior two thirds of body 5-6 txm, tapering to a sharply pointed tail. Sheath absent. Cuticle with fine transverse striations. Anterior end bluntly rounded. Inner body absent. Tail containing elongate nuclei, last nucleus not terminal. Fixed points expressed as percentage of body length as follows (one specimen from a thin smear of lung blood stained with Giemsa's): nerve ring 19; excretory vesicle 28; excretory cell 33; G1 64; anal vesicle 77.

Host: Corvus brachyrhynchos brachyrhynchos Brehm (CORVIDAE).

Location in host: Body-cavity.

Locality: Essex County, southern Ontario, Canada (winter flock).

Specimens: Parasitology Collection, National Museum of Canada No. NMCIC (p) 1979-1546.

List Ot species in the genus Cardiofilaria Strom, 1937

Cardiofilaria Strom, 1937 ( = Pseudaproctella Anderson, 1957; = Gallifilaria Jain, Alwar, Ad-

79


I i :....

b

t 7

2

t 8

80


Table III

Host distribution by avian family of species of Chandlerella Yorke and Maplestone, 1926

Body Location in Host: Heart Cavity Cavity

Avian Order ~ ~._ ~ = ~ - . ~ , ~ ~ ~ ~ .z ~0 = and Family ~-~ ~ "~. ~ ~ ~ ~ ~ ~ o

Connective tissue around blood vesseM: Other§

" ~ . ~ ~ ~ ' ~ ~ ~ - -

PELECANIFORMES Pelecanidae x Phalacrocoracidae

ACCIPITRIFORMES Falconidae

GALLIFORMES Tetraonidae Phasianidae

COLUMBIFORMES Columbidae

CAPRIMULG1FORMES Caprimulgidae

APODIFORMES Apodidae

PICIFORMES Rhamphastidae Picidae x

PASSERIFORMES Dicruridae Oriolidae Corvidae Certhidae Muscicapidae Motacillidae Sturnidae Icteridae Estrildidae

:x X

X X

X X

X X

X

X X

X

X X

X

X X X

* also reported from body cavity and connective tissue around blood vessels t also reported from heart cavity and connective tissue around blood vessels :~ includes locations reported as liver and kidney § C. braziliensis from cervical air sacs; C. quiscali from brain; C. petrowi from eye

wadhiya & Pandit, 1965; = Francofilaria Jairaj- puri & Siddiqi, 1970) contains 11 species. C. pavlovskyi is the generitype.

(i) C. pavlovskyi Strom, 1937 Synonyms: C. inornata (Anderson, 1956)

Sonin, 1961; C. ardae (Mawson, 1957) Sonin, 1961. Type host: Oriolus oriolus (Linnaeus) (ORIOLIDAE). Other hosts: ARDEIDAE: Ardea herodias

Fig. 1.

Fig. 2:

Fig. 3.

Fig. 4.

Figs. 1-8. Cardiofilaria pavlovskyi Strom, 1937

Cephalic extremity of female, apical view. Fig. 5. Posterior end of female, lateral view.

Anterior end of female, lateral view. Fig. 6. Posterior end of female, ventral view.

Microfilaria, lateral view. Fig. 7. Posterior end of male, ventral view.

Cephalic extremity of female, lateral view. Fig. 8. Posterior end of male, lateral view.

81


Table IV

Host distribution by avian family of species of Splendidolilaria Skrjabin, 1923

Location in Host: Heart Subcutaneous

+ - ~ . -,.b, . . . ~ ~ " ~ ' ~ . , ~ ~z

Avian Order ~ ~ ..~ ~ : , ' - ~ N ~ ~ ~ =~ '~ t - := = ~ -~ ~ ~ ~ ~ ~ ~ ~, and Family ~ ~ ' ~ " ~ ~ ~ ~ e ~ ~ . ~ ,. o . ~ = ~

Pul. art.~:

~ - , ' ~ ~

Other§

~ . , . ~ ' ~ . ~ ~..

G A V I I F O R M E S Gaviidae x

A N S E R I F O R M E S Anat idae x

A C C I P I T R I F O R M E S Falconidae

G A L L I F O R M E S Tetraonidae Phasianidae

C O L U M B I F O R M E S Columbidae

P S I T T A C I F O R M E S Psittacidae

C O R A C I F O R M E S Alcedinidae Meropidae

P ICIFORMES Ramphas t idae

P A S S E R I F O R M E S Tyrannidae Hirundinidae Corvidae Pycnonotidae Mimidae Muscicapidae Motacillidae Laniidae Sturnidae Nectariniidae Icteridae Fringillidae Ploceidae

X X X

X X

X

X

X X X X X X

X X X X

X

X X X

X X

X

X X

* also reported from the body cavity t also reported from the aorta :~ pulmonary artery § S. falconis f rom lungs; S. gretillati from body cavity; S. rotundicephala f rom eye; S. skrjabini f rom connective tissue

(Linnaeus), Botaurus lentiginosus (Mon- tagu) (see Mawson, 1957), Ixobrychus exilis (Gmelin) (see Gibson, 1973); ACCIPIT- RIDAE: Accipiter striatus velox (Wilson), Cirrus cyaneus hudsonius (Linnaeus) (see Anderson & Freeman, 1969); FAL- CONIDAE: Falco subbuteo Linnaeus (see Sonin, 1968); SCOLOPACIDAE: Philohela minor (Gmelin) (see Anderson &

Freeman, 1969); STRIGIDAE: Asio otus wilsonianus (Lesson) (see Anderson & Freeman, 1969); APODIDAE: Hirun- dapus caudacutus (Latham) (see Sonin, 1968); ORIOLIDAE: Oriolus chinensis Linnaeus (see Belopol'skaya, 1959, in Sonin, 1968); CORVIDAE: Corvus corax principalis Ridgway (see Anderson & Freeman, 1969); C. brachyrhynchos

82

Table V

Host distribution by avian family of species of Eufilaria Seurat, 1921"

Avian Order ~ ~ ~ "~" and Family ~ ~ N ~ ~ ~ ~ ~ ~

GALLIFORMES Phasianidae

GRUIFORMES Turnicidae

CUCULIFORMES Cuculidae

PASSERIFORMES Corvidae Campephagidae Pycnonotidae Muscicapidae Ploceidae Estrildidae

X

)4 :x[ X )(

X

X

X X

* All species have been reported from the subcutaneous tissues or the connective tissue surrounding the trachea and oesophagus.

(ii)

brachyrhynchos Brehm (present study); C. splendens Vieillot (see Wong, Poon & Anderson, 1980), Aphelocoma coerulescens (Bosc) (see Kinsella, 1974); MUS- CICAPIDAE: Hylocichla mustelina (Gme- lin) (see Anderson, 1956a), Monticola cinclorhynchus gularis (Swinhoe), Turdus dissimilis hortulorum Sclater (see Sonin, 1968), T. migratorius Linnaeus, Catharus ( = Hylocichla) ustulatus swainsoni (Tschudi) (see Anderson & Freeman, 1969), T. unicolor Tickell (see Majumdar, 1975), T. philomelos clarkei Hartert, T. iliacus Linnaeus (see L6pez-Caballero, 1977). MOTACILLIDAE: Anthus gustavi Swinhoe (see Sonin, 1968); LANIIDAE: Lanius cristatus Linnaeus (see Sonin, 1968); PARULIDAE: Geothlypis trichas (Linnaeus) (see Wells & Hunter, 1960), Seiurus aurocapiUus aurocapillus (Lin- naeus) (see Anderson & Freeman, 1969).

C. nuda (Hamann, 1940) Sonin, 1961 Type host: Chaetura pelagica (Linnaeus) (APODIDAE).

(iii)

(iv)

(v)

(vi)

(vii)

(viii)


C, dubia (Johnston & Mawson, 1940) Sonin, 1961 Type host: Geoffroyus (= Pseudopsittacus) geoffroyi maclennani (Macgillivray) (PSITI'ACIDAE).

C. graucalinum (Johnston & Mawson, 1940) Sonin, 1961 Type host: Coracina (= Graucalus) melanops (Latham) (CAMPEPHAGIDAE).

C. campanae (Chabaud & Golvan, 1956) Anderson & Freeman, 1969 Type host: Turdus iliacus Linnaeus (= T. musicus) (MUSCICAPIDAE). Other hosts: MUSCICAPIDAE: T. philomelus clarkei Hartert (see Chabaud & Golvan, 1956).

C. andersoni (Chabaud, Brygoo & Richard, 1964) Dissanaike & Fernando, 1965 Type host: Nectarinia souimanga apolis (Hartert) (NECTARINIIDAE).

C. major Dissanaike & Fernando, 1965 Synonyms: Pseudaproctella andersoni var. major Chabaud, Brygoo & Richard, 1964; C chabaudi Dissanaike & Fernando, 1965. Type host: Coua reynaudii Pucheran (CUCULIDAE). Comments: Dissanaike & Fernando (1965) regarded the specimens described as P. andersoni var. major by Chabaud et al. (1964) as a distinct species and proposed the name Cardofilaria chabaudi for it. However, the name major should have been raised to species status, according to the International Code of Zoological Nomenclature, Article 10b.

C. nilesi Dissanaike & Fernando, 1965 Type host: Gallus gallus Linnaeus (domestic chicken) (PHASIANIDAE). Comments: Infective larvae found in wild mosquitoes, Mansonia crassipes (Wulp), were given to chickens from which the type specimens were collected.

83


12

15 t

16

84

(ix)

(x)

(xi)

C. mhowensis (Jain, Alwar, Adwadhiya & Pandit, 1965) Anderson & Bain, 1976 Type host: Gallus gallus Linnaeus (domestic chicken) (PHASIANIDAE).

C. basiri (Jairajpuri & Siddiqi, 1970) An- derson & Bain, 1976 Type host: Francolinus pondicerianus (Gmelin) (PHASIANIDAE).

C. stepheni Schmidt & Neiland, 1973 Type host: Cymbilaimus lineatus fasciatus (Ridgway) (FORMICARIIDAE).

2. ChandlereUa chitwoodae Anderson, 1961 (Figs. 9-16, Table I)

Synonyms: Splendidofilaria [lexivaginalis Jones, 1961; C. flexivaginalis (Jones, 1961) Sultana, 1962; Parornitho]ilaria chitwoodae (Anderson, 1961) Sonin, 1965; P. flexivaginalis (Jones, 1961) Sonin, 1965; C. hispanica L6pez- Caballero, 1974.

General: Filarioidea, Onchocercidae, Splen- didofilariinae, ChandlereUa Yorke and Maples- tone, 1926. Loosely coiled, slender nematodes with attenuated extremities. Cuticle thin, generally smooth, but occasionally transversely striated near extremities. Cephalic extremity with four pairs of submedian papillae and lateral amphids. Oral opening small and elongated dor- soventrally. Buccal cavity absent. Oesophagus divided near nerve ring; muscular portion short, glandular portion longer and demarcated from intestine by slight constriction. Tail digitiform. Phasmids terminal.

Male (20 specimens): Length 10.7 (7.2- 13.6)mm. Maximum width 85 (70-110)~m,


near middle of body. Nerve ring 130 (100-167) m from anterior extremity. Muscular

oesophagus 140 (100-18(}) ~m long and 23 (18- 30) ~m wide. Glandular oesophagus 715 (580- 860) txm long and 32 (27-40) txm wide. Total length of oesophagus 855 (710-1030) ~m. Caudal extremity tightly coiled. Anus 101 (70- 130) txm from posterior end of body. SweLling of variable size present immediately anterior to anus. Four to five pairs of postanal papillae present in two rows on tail, often asymmetrical. Spicules subequal and slightly dissimilar, arcuate with pointed distal ends. Right spicule longer than left in 16 specimens, right spicule 100 (65- 124) tzm long, left spicule 84 (65-116) txm long. Tip of caudal extremity sometimes reflected dor- sally.

Female (20 specimens): Length 23.3 (16.0-27.8) mm. Maximum width 155 (107-207) txm, near middle of body. Nerve ring 143 (107-187) txm from anterior extremity. Muscular oesophagus 161 (127-190) tim long and 26 (16-30) ~m wide. Glandular oesophagus 827 (512-1013) ~m long and 33 (26-47) tim wide. Total length of oesophagus 988 (666-1233) tim. Vulva 414 (330-487) Izm from anterior end of body, slightly salient. Vagina 2100 (1100-2900) ~m long. Vagina usually extending short distance posteriorly, then bending anteriorly to pass vulvar region before turning posteriorly and joining uteri. Didelphic and opisthodelphic. Anus obs- cure, lateral in position. Tail 172 (107-260) ttm in length. Tip of tail sometimes knob-like.

Microfilaria (20 specimens from the blood): Length 190 (183-208) /xm. Width of anterior two thirds of body 5-6 tzm tapering to 4 txm in posterior third. Loose sheath present. Cuticle with fine transverse striations. Anterior end

Fig. 9.

Fig. 10.

Fig. l l .

Fig. 12.

Figs. 9-16. ChandlereUa chitwoodae Anderson, 1961

Anterior end of female, lateral view. Fig. 13. Posterior end of female, ventral view.

Cephalic extremity of female, apical view. Fig. 14. Posterior end of female, lateral view.

Microfilaria, lateral view. Fig. 15. Posterior end of male, lateral view.

Cephalic extremity of female, lateral view. Fig. 16. Posterior end of male, ventral view.

85

Filarioid nematodes of Cor~us brachyrhynchos brachyrhynchos Brehm

- ~ 1 8 20

17 ~"J 19

ol @

®

® ®

~J

I

25

86

bluntly rounded, cephalic hook and spines present. Inner body 15 to 25/xm long. Posterior end bluntly rounded, last nucleus terminal. Fixed points expressed as percentage of body length as follows: nerve ring 26 (24-27); excretory vesicle 37 (35-38); excretory cell 41 (38-42); anterior end of inner body 61 (58-63); posterior end of inner body 71 (69-74); G1 77 (72-81); R2 85 (83-87); R3 86 (84-89); Rn 88 (85-91); anal vesicle 89 (87-90).


Location in host: Connective tissue around vena cava, splenic artery, hepatic artery, mesenteric arteries, pulmonary arteries, brachial arteries and adrenal glands; subcutaneously in neck region.



3. ChandlereUa quiscali Robinson, 1971 (Figs. 17-25, Table I)

(von Linstow, 1904)

General: Filarioidea, Onchocercidae, Splen- didofilariinae, Chandlerella Yorke & Maples- tone, 1926. Slender nematodes with attenuated extremities. Cuticle delicate, smooth. Cephalic extremity with four pairs of submedian papillae and lateral amphids. Oral opening small and circular. Buccal cavity absent. Oesophagus in-


conspicuously divided in region of nerve ring. Two to six large cells surrounding oesophageal lumen immediately anterior to slightly con- stricted oesophago-intestinal junction. Caudal extremity bluntly rounded. Phasmids terminal.

Male (5 specimens): Length 25.7 (21.0-27.9) mm. Maximum width 120 (107-133) txm, near middle of body. Nerve ring 158 (150-170) ~m from anterior extremity. Muscular oesophagus 190 (150-230) txm long and 31 (30-33) txm wide. Glandular oesophagus 674 (650-730) ixm long and 51 (47-53) t.tm wide. Total length of oesophagus 864 (800-900) t~m. Caudal extremity tightly coiled. Tail 213 (180-250) txm in length. One pair of preanal papillae present or absent, when present often difficult to discern. Three pairs of postanal papillae present, in two rows on tail, often asymmetrical. Spicules subequal and slightly dissimilar, with rounded distal ends, right 73 (60-80) Ixm long, left 81 (73-90) txm long.

Female (10 specimens): Length 76.0 (62.1- 95.3) mm. Maximum width 164 (130-200) tzm, near middle of body. Nerve ring 170 (150- 200) tzm from anterior extremity. Muscular oesophagus 200 (170-230) txm long and 32 (30- 40) txm wide. Glandular oesophagus 850 (730- 1000) ~m long and 48 (40-50) tzm wide. Total length of oesophagus 1050 (930-1200) txm. Vulva 890 (770-1150) txm from anterior end of body, generally preoesophageal but near end of oesophagus, not salient. Vagina 1690 (1250- 2400) txm long, extending posteriorly. Wall in vulvar region composed of conspicuous cells. Di- delphic and opisthodelphic. Anus 440 (250-670) txm from posterior end of body.

Fig. 17.

Fig. 18.

Fig. 19.

Fig. 20.

Fig. 21.

Anterior end of female, lateral view.

Cephalic extremity of female, apical view.

Microfilaria, lateral view.

Cephalic extremity of female, lateral view.

Posterior end of male, ventral view.

Figs. 17-25. Chandlerella quiscali (von Linstow, 1904) Robinson, 1971.

Fig. 22. Posterior extremity of female, lateral view.

Fig. 23. Posterior end of male, lateral view.

Fig. 24. Posterior end of female, lateral view.

Fig. 25. Posterior end of female, ventral view.

87


Microfilaria (20 specimens, from the blood): Length 184 (178-193) ~m. Width of anterior two thirds of body 5-6/ tm tapering to 4 tzm in posterior third. Loose sheath present. Cuticle with fine transverse striations. Anterior end bluntly rounded, cephalic hook and spines present. Inner body 12 to 16/zm long. Posterior end bluntly rounded, last nucleus terminal. Fixed points expressed as percentage of body length as follows: nerve ring 25 (23-26); excretory vesicle 37 (35-40); excretory cell 41 (40-44); anterior end of inner body 62 (60-66); posterior end of inner body 69 (66-74); G~ 77 (75-82); R2 85 (83-88); R3 87 (85-89); R4 89 (85-91); anal vesicle 90 (88-92).


Vavilova, 1926, in Sonin, 1966); MUS- CICAPIDAE: Copsychus malabarica (Scopoli) (see Majumdar, 1975).

(ii) C. quiscali (von Linstow, 1904) Robin- son, 1971 Type host: Quiscalus quiscula versicolor Vieillot (ICTERIDAE). Other hosts: CORVIDAE: Corvus brachyrhynchos Brehm (see Jones, 1968 and present study), Cyanocitta cristata bromia Oberholser (see Cooper & Crites, 1974a); ICTERIDAE: Q. quiscula versicolor Vie- illot (see von Linstow, 1904, Odetoyinbo & Ulmer, 1959, Robinson, 1971, Granath & Huizinga, 1978 and Badley & Dronen, 1979).

Location in host: Beneath the pia mater of the cerebrum in the region of the sagittal fissure.



List of spedes in the genus Chandlerella Yorke & Maplestone, 1926

Chandlerella Yorke & Maplestone, 1926 (= Vagri]ilaria Augustine, 1937; = Skrjabinocta Chertkova, 1946; =Lerouxinema Singh, 1949; =Parornithofilaria Sonin, 1965) contains 23 species. C. bosei is the generitype.

(i) C. bosei (Chandler, 1924) Yorke & Ma- plestone, 1926 Type host: Dicrurus (= Dissemurus) paradiseus (Linnaeus) (DICRURIDAE). Other hosts: CORVIDAE: Corvus frugilegus Linnaeus (see Shikhobalova, 1933, in Sonin, 1966), Pica pica (Lin- naeus) (see Spassky & Oschmarin, 1939, in Sonin, 1966), C. splendens Vieillot (see Farooqui & Ahmed, 1977); CERTH- IDAE: Certhia familiaris Linnaeus (see

(iii)

(iv)

(v)

C. stantchinskyi Gilbert, 1930 Type host: Oriolus oriolus (Linnaeus) (ORIOLIDAE). Other hosts: CORVIDAE: Corvus corax Linnaeus (see Shikhobalova, 1933, in Sonin, 1966).

C. sinensis Li, 1933 Synonym: C. hawkingi Chatterjee, Sen & Bhattacharya, 1965 Type host: Dendrocitta [ormosae sinica Streseman (= Corvus sinensis), Urocissa erythrorhyncha (Boddaert) (= Urocissa sinensis ) (CORVIDAE). Other hosts: DICRURIDAE: Dicrurus forficatus (Linnaeus) (see Chabaud et al., 1964); CORVIDAE: Corvus frugilegus Linnaeus (see Shikhobalova, 1933, in Sonin, 1966), C. corone Linnaeus, (see Kurashvili, 1950, in Sonin, 1966), C. macrorhynchos Wagler (see Chatterjee et al., 1965); STURNIDAE: Acridotheres tristis (Linnaeus) (see Sonin & Bor- garenko, 1965, in Sonin, 1966).

C. columbigallinae (Augustine, 1937) Rasheed, 1960 Type host: Columbigallina passerina (Lin- naeus) (COLUMBIDAE).

88

(vi)

(vii)

(viii)

(ix)

(x)

(xi)

(xii)

(xiii)

(xiv)

(xv)

C. petrowi (Chertkova, 1946) Anderson & Freeman, 1969 Type host: Streptopelia orientalis meena (Sykes) (COLUMBIDAE).

C. lienalis Orloff, 1947 Type host: Alectoris gracea (Meisner) (PHASIANIDAE). Other hosts: PHASIANIDAE: Am- moperdix griseogularis iBrandt), (see Bor- garenko, 1960, in Sonin, 1966).

C. skr]abini (Petrov & Chertkova, 1947) Anderson, 1961 Type host: Falco subbuteo Linnaeus (FALCONIDAE).

C. lerouxi (Singh, 1949) Anderson & Pre- stwood, 1969 Type host: GaUoperdix spadicea (Gmelin) (PHASIANIDAE).

C. shaldybini Gubanov, 1954 Type host: Phalacrocorax urile (Gmelin) (PHALACROCORACIDAE).

C. pelecani (Chabaud & Rousselot, 1956) Anderson & Prestwood, 1969 Type host: Pelecanus ru[escens (Gmelin) (PELECANIDAE).

C. singhi Ali, 1956 Type host: Caprimulgus sp. (CAP- RIMULGIDAE). Other hosts: CORVIDAE: Urocissa erythrorhyncha (Boddaert) (see Ali, 1969).

(2. brasiliensis Yeh, 1957 Type host: Rhamphastos dicolorus Lin- naeus (RAMPHASTIDAE).

C. thapari Rasheed, 1960 Type host: Saxicola torquata (Blyth) (MUSCICAPIDAE).

indica

C. chitwoodae Anderson, 1961 Synonyms: C. flexivaginalis (Jones, 1961) Sultana, 1962 and C. hispanica L6pez-

(xvi)

(xvii)

(xviii)

(xix)

(xx)

(xxi)

(xxii)


Caballero, 1974 are herein designated as synonyms. Type host: Padda oryzivora (Linnaeus) (ESTRILDIDAE). Other hosts: TETRAONIDAE: Den- dragapus obscurus (Say), Bonasa umbellus (Linnaeus) (see Gibson, 1965); MUS- CICAPIDAE: Turdus iliacus Linnaeus, T. philomelos clarkei Hartert (see L6pez- Caballero, 1974); CORVIDAE: Corvus brachyrhynchos Brehm (see Jones, 1961, 1968).

C. buckleyi Sultana, 1962 Type host: Dendrocopas (=Leiopicus) mahrattensis aurocristatus (Tickell) (= Pi- cus blandfordi) (PICIDAE).

C. himalayansis Sultana, 1962 Type host: Falco (= Cerchneis) tinnunculus interstincms Horsfield (FAL- CONIDAE).

C. alii Sultana, 1962 Type host: Motacilla maderaspatensis Gmelin (MOTACILLIDAE).

C. apusi Sonin, 1963 Type host: Hirundapus (Latham) (APODIDAE).

caudacutus

C. striatospicula Hibler, 1964. Type host: Pica pica hudsonia (Sabine) (CORVIDAE).

C. columbae (Sonin, 1966) Anderson & Freeman, 1969 Synonym: (2. columbigallinae Rasheed, 1960 (not Augustine, 1937). Type host; Treron (= Crocopus) phoeni- coptera (Latham) (COLUMBIDAE).

C. longicaudata (Sonin, 1966) Anderson & Freeman, 1969 Synonym: C. sinensis Ali, 1956 (not Li, 1933) Type host: Corvus macrorhynchos Wagler (CORVIDAE).

89

Filarioid nematod e s of Corvus brachyrhynchos brachyrhynchos B reh m

27 3 0

I 2 6

I I • e 0

*1 0 e

Ii °

IP i - - e x v

--av /)

I ;O i

LN

~'.j 2 8 ,, [( ,, ,

29

~ PQ

o

Q

32

33

35

90

(xxiii) C. sultana (Sonin, 1966) Anderson & Freeman, 1969 Synonym: C. stantchinskyi Sultana, 1962 (not Gilbert, 1930) Type host: Lanchura (= Uroloncha) striata striata (Linnaeus) (ESTRILDIDAE).

4. Splendidofilaria caperata Hibler, 1964 (Figs. 26-35, Table I)

General: Filarioidea, Onchocercidae, Splen- didofilariinae, Splendidofilaria Skrjabin, 1923. Slender nematodes with attenuated extremities. Cuticle thick, with fine transverse striations and irregular, incomplete transverse thickenings and devoid of bosses. Cephalic extremity with two pairs of submedian papillae and lateral amphids. Oral opening small and circular. Buccal cavity absent. Oesophagus long and thin, not externally divided into muscular and glandular parts but occasionally increasing in width posteriorly. Oesophagus sometimes curved or looped before joining broad intestine. Caudal extremity digitiform, with two small terminal protuberances. Phasmids subterminal and ventrolateral.

Male (20 specimens): Length 13.7 (10.3-17.2) mm. Maximum width 136 (127-167) tzm, near middle of body. Nerve ring 127 (100-155) /xm from anterior extremity. Oesophagus 656 (390- 1020) ~m long and 11 (3-30) txm wide. Caudal extremity curved. Tail 99 (73-139) txm in length. Caudal papillae in two rows on tail, generally in pairs and often asymmetrical in arrangement. Two to four preanal, two or four adanal and four to five postanal papillae present. Spicules subequal and dissimilar, right 58 (51-70) /xm long with spatulate distal end, left 70 (60-80) tzm long with bluntly pointed distal end.


Female (20 specimens): Length 25.0 (12.7-36.3) mm. Maximum width 227 (150-290) txm, near middle of body. Nerve ring 138 (107-270) tzm from anterior extremity. Oesophagus 945 (590- 1700) txm long and 11 (5-25) ~m wide. Vulva 359 (280-570) gm from anterior end of body, slightly salient. Vagina 617 (307-1090)/xm long. Vagina often with one or two loops, rarely straight, occasionally extending anteriorly for a short distance before turning posteriorly and joining uteri. Didelphic and opistodelphic. Tail 105 (75-170) tzm in length.

Microfilaria (20 specimens from the vagina): Length 98 (85-120) /xm. Width of anterior two thirds of body 3-4/xm tapering to 2-3/~m in posterior third. Sheath absent. Cuticle with fine transverse striations. Anterior end slightly attenuated. Inner body present. Posterior end bluntly rounded, last nucleus terminal.


Location in Host: In the tunica media of the pulmonary arteries.



5. Splendidofilaria wehri Anderson, 1961 (Figs. 36-42, Table I)

General: Filarioidea, Onchocercidae, Splen- didofilariinae, Splendidofilaria Skrjabin, 1923.

Fig. 26.

Fig. 27.

Fig. 28.

Fig. 29.

Fig. 30.

Figs. 26-35. Splendidofilaria caperata Hibler, 1964.


Anterior extremity of female, apical view.


Cuticle, lateral view.

Anterior extremity of female, dorso-ventral view.


Fig. 32. Posterior end of male, ventral view.




91


36

38 39

41

Fig. 36.

Fig. 37.

Fig. 38.

Fig. 39.

I 5o ~m I

)

37

t o

Figs. 36-42. Splendidofilaria wehri Anderson, 1961

Anterior end of male, dorso-ventral view. Fig. 40. Posterior end of male, ventral view.

Oesophago-intestinal junction, dorso-ventral view. Fig. 41. Posterior end of male, lateral view.

Cuticle showing bosses, lateral view. Fig. 42. Posterior end of male, la[eral view.

Cuticle showing bosses, surface view.

Slender nematode with slightly at tenuated extremities. Cuticle thin, with fine transverse striations and transversely elongate bosses, except at extremities. Anter ior extremity with irregular transverse striations. Cephalic extremity with two pairs of submedian papillae and lateral amphids. Oral opening small and circular. Buccal

cavity absent. Oesophagus long and thin, increasing slightly in width posteriorly, not externally divided into muscular and glandular parts. Oesophago-intest inal junction clearly demarcated. Caudal extremity digitif0rm, with two small terminal protuberances. Phasmids not observed.

92

Male (one specimen): Length 8.5 mm. Maximum width 110/xm, near middle of body. Nerve ring 83/xm from anterior extremity. Oesophagus 390 txm long and 8-20 txm wide. Caudal extremity curved. Tail 70/xm in length. Caudal papillae grouped in anal region, two on right side, three on left. Spicules subequal, dissimilar, right spicule 45/xm long, distal end spatulate, left spicule 55/xm, distal end pointed.


Location in host: Behind one semilunar valve of the pulmonary artery.



List of species in the genus Splendido~laria Skr- jabin, 1923

Splendidofilaria Skrjabin, 1923 (= Ornithofilaria GiSnnert, 1937; = Ularofilaria Lubimov, 1946; = Parachandlerella Caballero, 1948; = Anenter- onema Oschmarin, 1949; = Lophortofilaria Wehr & Herman, 1956; = Splendidofilarioides Freitas & Ib~fiez, 1968) contains 28 species. S. pavlovskyi is the generitype.

(i) S. pavlovskyi Skrjabin, 1923 Type host: Lanius (= Otomela) collurio phoenicuroides (Schalow) (LANIIDAE). Other hosts: LANIIDAE: L. cristatus Linnaeus (see Sonin, 1966), L. (= Otomela) isabellinus Ehrenberg (see An- derson & Freeman, 1969); STUR- NIDAE: Sturnus (= Pastor) roseus (Lin- naeus) (see Sonin & Borgarenko, 1965, in Sonin, 1966); PLOCEIDAE: Passer domesticus (Linnaeus) (see Borgarenko, 1960, in Sonin, 1966).

(ii) S. mavis (Leiper, 1909) Anderson, 1961 Type host: Turdus iliacus Linnaeus (= T. musicus) (MUSCICAPIDAE).

(iii)

(iv)

iv)

(vi)

(vii)

(viii)

(ix)


Other hosts: M u s c i c A P I D A E : T. pilaris Linnaeus (see G6nnert, 1937), T. philomelos clarkei Hartert (see L6pez- Caballero, 1977).

S. gedoelsti Travassos, 1926 Type host: Selenidera maculirostris (Lich- tenstein) (RAMPHASTIDAE).

S. travassosi Koroliowa, 1926 Type host: Memps apiaster Linnaeus (MEROPIDAE). Other hosts: MEROPIDAE: M. super- ciliosus Linnaeus (see Sonin, 1961, in Sonin, 1966).

S. papillocerca (Lubimov, 1946) Ander- son & Chabaud, 1959 Type host: Tetraogallus altaicus (Gebler) (PHASIANIDAE). Other hosts: TETRAONIDAE: Tes- trastes bonasia (Linnaeus), Tet- rao urogallus Linnaeus, Lagopus lagopus (Linnaeus), L. leurcurus (Richard- son) (see Gibson, 1965); PHASIANI- DAE: Perdix barbara barbara Verreaux & Des Murs (= P. daurica), Tetraogallus altaicus (Gebler) (see Sonin, 1966).

S. periarterialis (Caballero, 1948) Chabaud & Choquet, 1953. Type host: Tyrannus sp. (TYRAN- NIDAE).

S. brevispiculum Singh, 1949 Type hosts: Streptopelia decaocto (Frivaldszky) (COLUMBIDAE); Hal- cyon smyrnensis (Linnaeus) (ALCED- INIDAE). Type host not indicated.

S. skrjabini (Oschmarin, 1949)Anderson and Bain, 1976 Type host: Garrulus glandarius (Lin- naeus) (CORVIDAE).

S. rotundicephala (Oschmarin, 1950) An- derson, 1961 Type host: Garrulus glandarius (Lin- naeus) (CORVIDAE).

93


(x)

(xi)

(xii)

(xiii)

(xiv)

S. verrucosa Oschmarin, 1950 Type host: Garrulus glandarius (Lin- naeus) (CORVIDAE). Other hosts: FRINGILLIDAE: Car- podacus erythrinus (Pallos) (= Erythrina erythrina) (see Sonin, 1963, in Sonin, 1966).

S. fallisensis (Anderson, 1954) Ander- son, 1961. Type host: Anas platyrhynchos Linnaeus (=A. boschas) (domestic Pekin duck) (ANATIDAE). Other hosts: ANATIDAE: Anas rubripes Brewster (see Anderson, 1956b), A. crecca Linnaeus (Daiga, 1965, in Sonin, 1966); GAVIDAE: Gavia immer (Briinnich) (see Anderson & Forrester, 1974).

S. algonquinensis (Anderson, 1955) An- derson, 1961 Synonym: S. passerina Koch & Huizinga, 1971 is regarded herein as a synonym of S. algonquinensis. Type host: Hirundo rustica erythrogaster Boddaert (HIRUNDINIDAE). Other hosts: HIRUNDINIDAE: Riparia riparia (Linnaeus) (see Sonin 1963, in Sonin, 1966); see PLOCEIDAE: Passer domesticus domesticus Linnaeus (see Koch & Huizinga, 1971); ICTERIDAE: Agelaius phoeniceus (Linnaeus) (see Cooper & (;rites, 1974b).

S. californiensis (Wehr & Herman, 1956) Anderson, 1961 Type host: Lophortyx californica californica (Shaw) (= L. californica vaUicola) (TETRAONIDAE).

S. tuvensis (Spassky & Sonin, 1957) An- derson, 1961 Type host: Perdix barbata turcom- ana Stolzman (=P. daurica) (PHASI- ANIDAE). Other hosts: TETRAONIDAE: Lagopus lagopus (Linnaeus), L. mutus (Montin), Lyrurus tetrix (Linnaeus), Tetra urogaUus

(xv)

(xvi)

(xvii)

(xviii)

(xix)

(xx)

(xxi)

(xxii)

Linnaeus, Tetrastes bonasia (Linnaeus), Falcipennis falcipennis (Hartlaub) (see Sonin, 1966).

S. bohmi (Supperer, 1958) Anderson, 1961. Type host: Turdus viscivorus Linnaeus (MUSCICAPIDAE).

S. gretillati Chabaud, Anderson & Brygoo, 1959 Type host: Agapornis cana (Gmelin) (PSITrACIDAE).

S. kashmirensis Amir & Ali, 1960 Type host: Pycnonotus (= Molpastes leucogenys leucogenys (Gray) (PYCNO- NOTIDAE).

S. wehri Anderson, 1961 Type host: Catharus (= Hylocichla) ustulata swainsoni (Tschudi) (MUSCI- CAPIDAE). Other hosts: CORVIDAE: Corvus brachyrhynchos brachyrhynchos Brehm (present study).

S. singhi Sultana, 1962 Type host: Motacilla maderaspatensis Gmelin (MOTACILLIDAE).

S. alii Sultana, 1962 Type host: Nectarinia ( = Leptocoma ) asiatica asiatica (Latham) (NECTARIN- IIDAE). Other hosts: STURNIDAE: Sturnus (=Pastor) roseus (Linnaeus) (see Ali, 1969).

S, picacardina Hibler, 1964 Type host: Pica pica hudsonia (Sabine) (CORVIDAE).

S. caperata Hibler, 1964 Type host: Pica pica hudsonia (Sabine) (CORVIDAE). Other hosts: STURNIDAE: Sturnus vulgaris vulgaris Linnaeus (see Ballard, 1965 and Ballard & Olsen, 1966); COR-

94

(xxiii)

(xxiv)

(xxv)

(xxvi)

(xxvii)

(xxviii)

VIDAE: Corvus brachyrhynchos brach- rhynchos Brehm (present study).

S. falconis (Sonin, 1966) Anderson & Freeman, 1969 Synonym: Chandlerella bosei Koroliowa, 1926 (not Chandler, 1924). Type host: Falco tinnunculus Linnaeus (FALCONIDAE).

S. pectoralis Gibson, 1967 Type host: Dendragus obscurus pallidus Swarth (TETRAONIDAE). Other hosts: TETRAONIDAE: Canachites canadensis (Linnaeus), Bon- asa umbellus (Linnaeus), Pedioecetes phasianellus (Linnaeus) (see Gibson, 1967).

S. pachacuteci (Freitas & Ibfifiez, 1968) n. comb. Type host: Mimus longicaudatus Tschudi (MIMIDAE).

S. longicaudata Devamma, 1977 Type host: Chrysomma sinene (Gmelin) (MUSCICAPIDAE).

S. osmaniae Devamma, 1977 Type host: Sturnus (= Temenuchus ) pagodarum (Gmelin) (STURNIDAE).

S. gvozdevi Sonin & Barus, 1978 Type host: Alectoris graeca (Meisner) (PHASIANIDAE).

6. Eufilaria longicaudata Hibler, 1964 (Figs. 43-50, Table I

General: Filarioidea, Onchocercidae, Lem- daninae, Eufilaria Seurat, 1921. Small, slender nematodes with bluntly rounded extremities. Cuticle thin, smooth, with fine transverse striations and occasionally with larger transverse striations near anterior extremity. Cephalic extremity with four pairs of submedian papillae and minute lateral amphids. Mouth opening small and circular. Buccal cavity absent. Oesophagus


short and thin, not externally divided into muscular and glandular parts; indistinctly separated from intestine.

Male (7 specimens): Length 8.1 (5.8-10.6) mm. Maximum width 64 (40-90) /xm, near middle of body. Nerve ring 158 (115-175) txm from anterior extremity. Oesophagus 311 (200-450)/xm long, 7 (8-15) fzm wide. Caudal extremity straight. Anus 13 (10-20) ~m from posterior end of body. One large lateral swelling present on both sides of tail slightly behind anus; zero to four minute papillae present on each swelling. Spicules subequal and dissimilar, right 59 (47- 82) txm long with pointed distal end which in ventral view appears arrow-shaped, left 67 (53- 85) ~m long with pointed distal end. Phasmids subterminal.

Female (3 specimens): Length 16.6 (13.7-18.0) mm. Maximum width 128 (95-160) ttm, near middle of body. Nerve ring 175 (160-200) ttm from anterior extremity. Oesophagus 258 (195- 310) ~m long, 13 (7-22) ttm wide. Vulva 552 (450-680) ttm from anterior end of body, slightly salient. Vagina 893 (450-1150)/xm long, directed posteriorly or extending anteriorly before turning posteriorly. Didelphic and opisthodelphic. Anus 46 (32-67) ttm from posterior end of body.

Microfilaria (20 specimens from the blood): Length 108 (95-123) ~m. Width of anterior two thirds of body 5-6txm, tapering to a sharply pointed tail. Sheath absent. Cuticle with fine transverse striations. Anterior end bluntly rounded, oral ring and spines present. Inner body 4 to 7/~m long. Tail containing three or four elongate nuclei, last nucleus not terminal. Fixed points expressed as percentage of body length as follows: nerve ring 22 (17-25); excretory vesicle 33 (29-37); excretory cell 37 (35-41); anterior end of inner body 56 (49-60); posterior end of inner body 61 (55-65); G1 66 (61- 73); R2 73 (67-79); R3 75 (68-82); R4 77 (70-84); anal vesicle 80 (74-86).


95


Fig. 43.

Fig. 44.

Fig. 45.

Fig. 46.

i

J

45

B

Figs. 43-50.


Anterior extremity of female, lateral view.

Anterior extremity of female, apical view.


o o , .'

i ~ m,',r'~L'

47

J

Eufilaria longicaudata Hibler, 1964





Location in Host: Connective tissue around oesophagus and trachea.

List of species in the genus Eufilaria Seurat, 1921



Eufilaria Seurat, 1921 (=Eufilariella Sonin, 1965; =Neofilaria Deshmukh, 1968) contains 10 species. E. sergenti is the generitype.

(i) E. sergenti Seurat, 1921

96

(ii)

(iii)

(iv)

Type host: Passer hispaniolensis (Tem- minck) (PLOCEIDAE). Other hosts: CORVIDAE: Garrulus leucolophus* (see Yeh, 1957).

E. capsulata (Annett, Dutton & Elliot, 1901) Seurat, 1921 Type hosts: Pycnonotus barbatus (Des- fontaines) (PYCNONOTIDAE): Ploceus (=Sitagra) nigricollis brachypterus Swain- son (PLOCEIDAE); Ploce us (= Hyphantor- nis) sp. (PLOCEIDAE). Type host not indicated.

E. asiatica Singh, 1949 Type host: Corvus splendens (CORVIDAE).

(Vieillot)

E. delicata Supperer, 1958 Type hosts: Turdus viscivorus Linnaeus (MUSCICAPIDAE); T. merula Linnaeus (MUSCICAPIDAE); Garrulus glandarius (Linnaeus) (CORVIDAE). Type host not indicated.

(v)

(vi)

(vii)

(viii)

(ix)

U. mcintoshi Anderson & Bennett, 1960 Type host: Padda oryzivora (Linnaeus) (ESTRILDIDAE).

U. singhi (Sultana, 1961) Anderson & Pre- stwood, 1969 Type host: Coracina (= Lalage) melanopt- era sykesi (Strikland) (CAMP- EPHAGIDAE).

E. Iongicaudata Hibler, 1964 Type host: Pica pica hudsonia (Sabine) (CORVIDAE). Other hosts: CORVIDAE: Corvus brachyrhynchos brachyrhynchos Brehm (present study).

E. alii (Deshmukh, 1968) Anderson & Bain, 1976 Type host: Turnix tanki Blyth (TUR- NICIDAE).

E. buckleyi (Deshmukh, 1968) n. comb. Synonyms: Neofilaria buckleyi Deshmukh,

* no t l is ted in Pe t e r s ' Check- l i s t of Bi rds of the W o r l d


1968; E. utae Anderson & Bain, 1976. Type host: Perdicula asiatica (Latham) (PHASIANIDAE).

(x) E. coua Anderson & Prestwood, 1969 Type host: Coua reynaudii Pucheron (CUCULIDAE).

Note: E. cypseli (Annett, Dutton & Elliot, 1901) Nelson, 1966 does not belong in Eufilaria. The species is incompletely described and it is difficult to ascertain the genus in which it should be placed. However, because the spicules are unequal in length, it is transferred provisionally to Lemdana as L. cypseIi (Annett, Dutton & Elliot, 1901) n. comb. Sonin (1966) placed E. buckieyi Rasheed, 1960 as a synonym of Aprocta obtusa (Dujardin, 1945) Sonin, 1966.

Discussion

Chandlerella chitwoodae was the most common filarioid parasitizing crows in the present study. Jones (1968) found C. chitwoodae (= C. flexivaginalis) in 33% of 339 crows collected in OhiO. Robinson (1955) found a microfilaria, the brief description of which fits that of the microfilaria of C. chitwoodae, in 65% of 217 crows collected in Georgia. Thus, C. chitwoodae is probably a common parasite of crows throughout eastern North America. In addition, Robinson (1955) found a similar microfilaria in 1.6% of 490 blue jays (Cyanocitta cristata) collected in Georgia. C. chitwoodae has also been reported in mannikins (Padda oryzivora) by Anderson (1961a), in grouse (Dendragapus obscurus and Bonasa umbellus) by Gibson (1965)** and in thrushes (Turdus iliacus, T. philomelos) by L6pez-Caballero (1974)***.

Splendidofilaria caperata was the second most common filarioid encountered. Microfilariae of S. caperata, however, were never found in the peripheral blood of infected crows and infection with this species is regarded as occult. Uniden- tified adult filarioids found by Elliot (1902) in

** Gibson (1965) iden t i f i ed C chitwoodae as C. flexivaginalis. *** L 6 p e z - C a b a l l e r o (1974) ident i f ied C. chitwoodae as C. hispanica.

97


the walls of the pulmonary arteries of crows collected in Ontario were possibly S. caperata. This species was reported in 41% of 124 magpies (Pica pica hudsonia) by Hibler (1963) and in 3% of 70 starlings (Sturnus vulgaris) by Ballard (1965), both hosts having been collected in Col- orado, USA.

Eufilaria longicaudata was slightly less common than S. caperata in crows. Robinson (1955) reported an unsheathed microfilaria with a sharply pointed tail in 13% of 217 crows and 55% of 490 blue jays collected in Georgia. His description agrees with that of the microfilaria of E. longicaudata and it may well have been this species, although precise identification is impos- sible. Hibler (1963) found E. longicaudata in 59% of 142 magpies collected in Colorado.

Chandlerella quiscali was found in only 1.9% of the crows examined. In Ohio, C. quiscali occurred in 14% of 339 crows collected by Jones (1968) and in 2% of 50 blue jays collected by Cooper & Crites (1974a). This species was also reported in 62% of 112 grackles collected in Iowa (Odetoyinbo, 1960), all of 88 in Illinois (Granath & Huizinga, 1978) and 57% of 21 in Texas (Badley & Dronen, 1979).

Only 0.4% of 269 crows examined were infected with Cardiofilaria pavlovskyi. This species is readily observed in the abdominal cavity of an infected bird and its repeated absence from helminth surveys of the crow (Ward, 1934; Morgan & Waller, 1941; Daly, 1959; Jones, 1968, Hen- dricks et al., 1969; Andrews & Threlfall, 975) is considered significant. C. pavlovskyi is known from 26 additional species of birds in 13 families.

Splendidofilaria wehri occurred in only 0.3% of the crows examined in the present study. It has also been reported in the olive-backed thrush ( Catharus (= Hylocichla) ustulata swainsoni) in Algonquin Park (Anderson, 1961b).

Chandlerella chitwoodae, S. caperata and E. longicaudata occurred commonly in the present study and are considered enzootic in the crow population. Both C. chitwoodae and E. longicaudata produced a microfilaraemia in crows. S. caperata produced occult infections and must be commonly acquired from other hosts in the environment of the crow. C. quiscali, C. pavlovskyi and S. wehri occurred infrequently and are

undoubtedly sporadic infections in the crow population. They also are probably acquired from other birds. For example, C. quiscali is a common parasite of grackles and may spread to crows from this host.

Although more than 40 genera have, at one time or another, been proposed to accomodate avian filarioids, only 15 genera are currently recognized (Anderson & Bain, 1976). This raises the possibility that only a few avian filarioid genera exist, most of which are now known. Cardiofilaria, Chandlerella, Splendidofilaria and Eufilaria are four of the largest and most clearly defined of these genera.

Species of these four genera have been reported from numerous families of birds in different orders (Table II to V). Thus, they are not restricted to particular taxonomic groups of birds. In addition, some species, for example Chandlerella chitwoodae and Cardiofilaria pavlovskyi, are apparently not highly host specific. Further study may reveal that this is also true for other species. Those species which are not highly host specific probably parasitize groups of birds living in similar habitats and in association with certain vectors.

Filarioids with a broad host distribution are probably transmitted by vectors with non-specific feeding habits. Microfilariae of Chandlerella chitwoodae, for example, will develop to the third stage in Culicoides stilobezzioides Foote & Pratt and in C. travisi Vargas (Bartlett & Ander- son, in press). Both midges are woodland in- habitants and appear to have non-specific feeding habits (Bennett, 1960; Fallis & Bennett, 1960, 1961; Snow, 1955; Pickard & Snow, 1955; Robinson, 1971). Thus, Chandlerella chitwoodae may be commonly transmitted among crows and other birds living together in a woodland habitat which includes such vectors.

The vectors of Cardiofilaria pavlovskyi are not known, but they probably also have non-specific feeding habits. C. nilesi is transmitted by the mosquito Mansonia crassipes (Wulp) (see Niles et al., 1965; Dissanaike & Fernando, 1965). Mosquitoes may, as suggested by Anderson & Freeman (1969), be involved in the transmission of C. pavlovskyi. Mosquitoes in the genus Culex are, in general, nonspecific bird feeders (James &

98

Hardwood, 1969) and might transmit avian filarioids in addition to avian malaria and viruses.

A filarioid which is not host specific probably parasitizes a number of hosts within a specific bird community but one host species may be more important than others in maintaining the infection in the community. This main host species may differ from one region to another. For example, the woodcock is apparently the main host of Cardiol'ilaria pavlovskyi in Ontario (Anderson & Freeman, 1969) but, in Florida, the scrub jay is possibly more commonly infected (Kinseila, 1974).

A filarioid transmitted by a vector which feeds in a specific stratum of a woodland habitat would probably occur with different levels of prevalence in birds tending to occupy different parts of the habitat. For example, Bennett (1960) collected Culicoides stilobezzioides from a variety of birds exposed above ground level in a woodland community but not from those exposed at ground level. If Chandlerella chitwoodae were transmitted solely by this midge, it would probably be common in birds living above ground level but sporadic in those living at ground level.

The genera Cardiofilaria, ChandlereUa, Splen- didolilaria and Eulilaria are not restricted to any particular zoogeographical regions and some species, e.g. Cardio1'ilaria pavlovskyi and Chand- lerella chitwoodae, apparently have a cosmopoli- tan distribution. Such a wide distribution is not surprising because birds are capable of flight and assuming the presence of suitable vectors, may spread infections to susceptible bird populations in different geographical regions. It is possible that migrating birds which stray from their usual migration routes introduce infections into new areas and thus play key roles in the world-wide dissemination of filarioid species.

Acknowledgements

Sincere thanks are extended to J. Branch, R. Jackson, M. Leonard and P.L. Wong for invalu- able assistance in the collection and dissection of crows. The advice of Mrs. U. Strelive in the preparation of drawings is gratefully acknow- ledged. Mr. W.O. Granath (Illinois State Univer-


sity), Dr. D.C. Gomes (Instituto Oswaldo Cruz), Dr. C.P. Hibler (Colorado State University), Dr. J.R. Lichtenfels (US National Museum) and Dr. E.J. L6pez-Caballero (Universidad de Granada) kindly lent specimens. This study was supported by a NSERC of Canada operating grant to R.C. Anderson and a postgraduate scholarship to C. Bartlett.

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Accepted for publication 16th January, 1980.

Addendum: Since this paper went to press the following article has come to our attention: Olsen, O.W. & Braun, C.E. (1976) New species of Splendidofilaria and Chand- lerella (Filarioidea: Nematoda), with keys to the species, from the band-tailed pigeon (Columba fasciata fasciata) in the Rocky Mountain region. Great Basin Naturalist, 36, 445-457. In their paper the authors describe the new species Splendidofilaria columbensis, S. hibleri and Chand- lereUa robinsoni. S. columbensis and S. hibleri are known from males only and their validity requires further study. C. robinsoni will also require further study.

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