439

PART Section C. _Z

Phylum ASCH) LMINTHSS

Class Nematoda. 440

PHYLUM ASCHELÄINTHES Grobben, 1910

Syns.: 2lemathelminthes Gegenbaur, 1859 in part ; Vematoda ([ (Rudolphi, 1808) Diesing,. 1861 Lankester. ' 1477) Potts, `-1932t,,,

Nemata [ (Rudolphi, 1808) Cobb, 1919 ], Pearse, 1936; Nemerta

([Oersted,. 1844 ] Lankester, - 1900) Chitwood, 1950. '

Class NEMATODA (Rudolphi, 1808) einend. Diesing, 1861-, * & Chitwood, 1933) 1'earse, Syns.: Aphasraidia -(Chitwood -1936; '- Phasmidia (Chitwood and Chitwood, ' 1933) Pearse, 1936;: Adenophorea. (von Linstow, 1905) Chitwood,, 1958 (b); Secernentea (von Linstow,,

1905) Chitwood, '1958 (b)" E(von Linstow, 1905) Chitwood, 1940 nom., Sub-class Adenophoria . d, ', 1933; e; nend. Syns.: Aphasmidi a- Chitwoo d and, Chitwoo '.

Adenophori, (von Linstow, 1905) Chitwood, 1940.

Sub-class Secernentia ((von Linstow, 1905) Chitwood, ly4U/ nom.

emend. Syns.: Phasmidia Chitwood & Chitwood, 1933;

Secernentes`(von Linstow, -1905) Chitwo'od, ý, 1940., S While descriptions of new species and genera of free-living ', and zoo- and phyto- parasitic nematodes continue'to be published', increasing the classification. of the, group remains ---' , in numbers, to zoological thought., It ' is -precisely because an, obstacle all, _ , , ,-' she is able to bring the clear-eight of-a-zoologist rather-than`-. near-sight nematologist the writer has -followed ,,, the of a -that (1951) in using Aschelminthes'as the name. --, - Dr. Hyman phylum --The; phylum Nemathelminthes-has been utilised-to include'the acantho- of cephalans - but there are valid reasons for the ' exclusion these ,'ti of animals from, a' common phylum with the' nematodes',. not ° least - which is the fact that much, of their internal organization, shows.,:

a closer affinity to the cestodes, phylum platyhelminthes. ', As

understood, the Aschelminthes contain six classes; the'Rotifera, -,

Gastrotricha, Echinodera, -Priapulida, Itematoda and Nematomorpha

"with'_the following. definition: - pseudocoelomate with a'cuticle. =

and hypo-cuticle'secreted by the ecto dermal cells and a. -lbasically

ground planýfor the Steiner (1960), similar excretory system. , has mentioned that Nemathelminthes should have precedence over

'Aschelminthes because of priority, but, this phylum name,., unless'_`. -

specifically emended, must be-understood to include the Acantho-

cephala. a) Chitwood (1958. withdrew his sub-classes Aphasmidia and,. '_

Phasmidia, expressing regret that he had not acted on-his,, l940`.

acceptance of Adenophori and Secernentes as correct. 'He, left,,

these names open for usage as either-sub-class or class, Pearce, '

(1936) having already promoted the, classes Phasmidia'and' (1958) Aphasiaidia. Dougherty, ' accepted Phylum , Nematoda. Later

that year in the'discussion that followed Chitwood's contribution

to the International Congress for Zoologyý(1958), Dougherty

stated that he considers Aschelminthes to be`the valid phylum'

name and, Nenatoda'a class. 'This-relegation to class rankeis well (1960): '"To'consider_the -expressed by Steiner Nematoda aphylum',:.

would make it'necessary'to, elevate also, most, if, not'all, of the,

related taxa to phylum rank, a''proceeding that would signify a, '

" negation : of an existing and ,.unquestionable relationship" .; if :.,

the classes -Adenophorea. and Secernentea `are to- be' reduced- to .`. sub-classes it-is necessary to e.aend their terminations to the,

accepted 'ia'.,.

The classification of Yamaguti (1961) consists re- . of a woricing of Baylis and Daubney (1926) which, consistent for . while, zoo- nematodes,., ignores both free-living parasitic . and phyto-, parasitic nematodes. The entire group is so-remarkable in-its . morphological uniformity as opposed, to the-'biological, `physiol- -:

ogical'and ecological adaptations that from a purely zoological .- of point view a discipline which'attempts to, embrace this variety

is to be preferred. The general dissatisfaction -with- the' presence

of absiance of phasmids as a criterion arid the reluctant acceptance

of, the Chitwood classification as given in Thorne (1961)_p. 89,,.

does not negate the necessity for a system which places the

natodes in their broadest context. '. Only Dr Chitwood ' is ne; , o' ,;: qualified and has attempted to do-'s o and ' so must receive `due

honour.

In an attempt to determine the claesifi'eation acceptable: to

other workers the writer has. consulted the last four taxononjc

theses submitted, from this Department. Khalil(1962) and-Wahid,

ý,(1962). both refer only to families and. neither in their intro--

duätions state. the classifications utilised. Tadros '(1965)

adheres rigidly to Yamaguti (1961)., wibhout reference to -Chitwood..

This general, reluctance to ' discuss the merits of,: or reasons, for ., rejecting, '> the various, systems of higher, classification is, found . in Rasheed, (1964). This last also - worker, whose knowledge of5

:, the class is very. sound and who'. might have been expected- to 443 record reasoned views on general classification, makes no attempt to include a classification beyond Super-families - an omission which must have been intentional.

If taxonomic theses are so circumscribed in outlook as to ignore general zoological implications, while adding to the and profusion confusion of generic, specific and sub-specific names, it is only to be expected that taxonomy based on morphology is being ridiculed and the claims of the computer endorsed. The writer accepts the more recent scheme of Chitwood (1959 - quoted

Thorne, 1961) at subclass level as zoologically correct.

<: 444 Order: DORYLAIMIDA Pearse, 1936

Super-family: -TRICHUROIDEA Railliet, 1916

Family: 'Trichuridae Railliet, 1915

Capillariinae Railliet, Sub-family: 1915: ý

Railliet (1916) recognised, two families, Trichinellidae'and'

Trichuridae, the latter possessing Trichurinae,, Capillariinae, and

Trichosomoidinae. Chitwood (1937) and Chitwood and Chitwood (1950) have Railliet's In Neveu- -, accepted, classification. - , '1936, Lemaire created -a separate family for Capillaria', and this has-

oeen'adopted'by, Skrjabin, Shikhabolova and Orlov (1957).

Genus: - Capillaria Zeder, ý 1800. Syns.: Gordius Goeze, 1782; Trichice halus_Schrank, '1788;

Rudolphs, 1819; Trichosomu: n Creplin, Lin iscus ''Trichosoma -1829; '_ Duj ardin, 1845; Thominx Duj ardin, 1845; Eucoleus Duj ardin, 1845;

Ca1odium Duj ardin, 1845; Hepati cola= Hall, 1916; ` Capilloetronxv_ i'looides Freitas and Lent, 1935; Aonchotheca, Lopez-Neyra, 1947;

Echinocoleus Lopez-Nbyra, 1947. '

Goeze (1782) described-Gordius from hens, and Schrank (1788)

described Filaria gallinae and in 1790,. -Trichicephalus anatis. F. gallinae to ' be Gordius and are now -considered synonyms, -of C. caudinflata and T. anatis a synonym`of, C. anatis. Aadaen, '1951. (1800) described - Zeder, Capillaria : but , this name was displaced by Trichosoma of Rudolphs (1819). Dujardin, '(1845) 'separated, genera as five follows: -

I). -Sheath short,, -equal to twice the'diameter of,, the body, ' smooth not thickened: - Tricho'soma'(_'Capillaria) .445 : II). Sheath large, ten times the diameter of the body with spines:.

(a) spicule present -'Thominx Eucoleus _(b) spicule absent" - III). Sheath very large, no spines but ' with- transverse, - oblique

striations not thickened, 'and frequently projecting: -' Cal=, Liniscus.

Diesing'(1851) recognised only one genera but several sub-.. (no spicule ' genera: Gymnothecae spines on - sheath) - Calodiua,, Echinothecae (seined, Thominx, 'Eucoleus Liniscus;, spicule -sheath)' --: In his 1861`; revision he accepted all genera but Lin iscus.

Stossich (1890) added a, third'sub-genus, Athecae"for', Trichosoma-'.

has The Zeder's-- crassicauda which , -no sheath. priority of ._, Capillaria was first- acknowledged by Travassos (1914) who, synon-', -'

this The following -ymised all-other genera under name. year created two sub-genera: -,

Capillaria, (C. )'-, Zeder, , 1800 '- non-spinose spicule 'sheath

Capillaria (Thominx) Dujardin, 1845 -ýspined spicule. sheath.

The genus Hepaticola was raised, by Hall (1916).. for'eo 8ca from the liver of mammals, but `Baylis (1931) concluded, that _ , most diagnostic characters were of negative - value and: synonymised ýall-,

with ' Capillaria Zeder, . 1800. His decision has., been, genera _ -, followed by Cram, (1936) 'and Ldadcen (1945), the' latter's revision

of certain avian species being extensive. Skrjabin (1939);

Thoainx Dujardin, 1845, the, re-established . with -thespines on, sheath rating generic , importance ` and'in addition to ; accepting,

this work, Skarbilovitsch (1946)' proposed Skriabinocapillari for ` species in bats.

Lopez-Neyra (1947) based a new classification on the ratios of oesophagus length to'that of the body and the presence or absence of spines on the spicule sheath, but Madsen (1,95l)-showed that this scheme is unreliable and reverted to Baylis'. ', classif- -', , ication. Gagarin (1951) stuaied avian species and conceded three genera with the hitherto=unconsidered feature of positionAn-the =-viz., `, host utilised as well as morphological characteristics, _ of Eucoleus -'mucosa of oesophagus, Capillaria - lumen small, intestine, Thominx - within the caecum. - Skriabinocapillaria was "and added to this classification, by`Chetkovä (1952) Hepaticola' was considered valid. -by Pavlov (1955) because of the unusual site in. the host and the unique life-cycle.

The next majPr classification was contributed by Skrj abin, ` .' which Shikhabolova and Orlov in 1957, iaay sut marised as follows: _be' , 1) Spicule present .. 2, ...... 5.. -.. .. -.. .., .. Spicule .... :. `, '. ,... absent '`.. .. . 5 ..., 4- 2) Spicule .... sheath with'spines .. "-.. .. Thominx Spicule sheath without spines 061 41."...... by female from 3) Eggs-laid emerge the host .. Capillarig Eggs laid by female remain within the host, '... Hepaticola ý` _, , , Spicule Skriabinocapill 4) sheath without spines, . .... ria Spicule ° '- Euco1eý : sheath with spines-! ...... ,..: "ý _.. .. _ Capillostrongyloides Preitas and Lent, 1935 and, Aonchotheea Lopez-Neyra, 1947, Capillaria, ` Echinocoleusý.. are placed under and', ',. Lopez-Neyra, 1947, under Thominx. A curious feature of: this is the C. 1945 -extensive revision nomination of obsignata Madsen, as genotype instead of. C. columbae (Rud., 1819). Neither

columbae not obsignata can justifiably replace anatis (Schrank, ,-. 1790) the by Even as genotype accepted nearly, all . other workers. follow workers who this classification express doubts similar to (1931), Stewart (1963) in those of Baylis -for example T, Bonner,

a note'on-Thominx phasianina (Kotlan, 1940) states "because of the lack of cuticularization of the spicule of certain capillarids,

the presence or absence of'a spicule is not easily determined". `

One suspects that a character which may or may not be-. -presentý

in species should not rank at a. generic level. From within_` 'a .:: comes the Soviet Union the'criticism of Ga'garin, (1959)'-that' the

division of Capillaria into genera has been done without a

thorough study and redescription of species from all-groups of, -

hosts., -

Freitas (1959) added four new, genera, Gessvella, 'Pterothomini,

Pseudocapillaria and Ritaklossia, using'the: criteria, - nature of 'relative `lengths the cephalic end, of anterior-, and, posterior-,.. _, mature regions, nature of the eggs and of, the'aale, caudalýregion.

Freit4s ' and, I[endonca (1960) created Pear'soneina;. for some mammalian _ specigs,, jind Freitas and Silva, (1960) Orthothominx for, various's.

avian'slecies. '_Yamaguti (1961) has created'a new Order

Trichiirtdea (syn., Trichinelloidea, Hall, ``1916), while the Chitwoods (1950) place 'the family in' Dorylaimida , Fearse, 1936.

Yamaguti rejects- Skrjabin' et: ' al., '(1957) -and 'synonymises' all, species in - Capillari a with the, exception, of Skri abinocapillaria ,. 448 which he separates with a footnote; to the effect that the poorly chitinised spicule may have been overlooked.

In this-study all specimens are considered as belonging to-,

Capillaria in agreement with_Baylis, (1931),, Cram_(1936), Madsen;

(1945, Read (1949) and the `, Chitwoods, (1950). ý t-19519,1952)1, .

Capillaria (Molin, 1858) Wavilova, 1926 '- 1). caudinflata - ý Sins".: Gordius gallinae Goeze, - 1782; , Filaria' aallinae -_-, Schrank, 1788; Capillaria semiteres 'Zeder, ` 1803; 'Trichosoma

Rudolphi, `1819; Trichosoma' longicolle Rudolphi,, ".1819; columbae - Calodium tenue Duj ardin, 1845';, Trichosoma gallinum Kowalewaki,

1894; C., longicollis, Madsen, 1945; C. gallinae Lopez-NAyra, '1947;

Thominx wavilovi Skrjabin et al., 1957 and many others.

The first accurate and-recognisable description of-this

species is that of, Aplin (1858) as -Calodium-"caudiznflata from the. , quail (Coturnix, coturnix (L. )) and £adsen (1951) conceded that.: result his longicollis of 1945 is "a- synonym- as a of the work ; of

Orosz (1931> and Moorehouse (1944). ' Molin '(1859)did not . -'mention the presence, of spines-on the spicule sheath and despite their-

presence in Madsen's redescription (1945), Skrjäbinet al`. (1957) longicollis, from separated caudinflata on the -presence of 'spines,,

and, used it for their new 'species Thominx wavilova. °-"Freital_

(1960), used, the presence of spines and-lateral-caudal alae°. to BLANK IN ORIGINAL platet 42

Capillaria caudinflata (golin, 1858) Wavilova, 192b

Figure 135 Male Tail. Bursa lateral view. H. P., 1 10 Figure 136 Male'Tail. Bursa ventral view. N. P. I IO Figure 137 peXale. 'Vulva appendage. N. P. x 10. Figure 138 Egg. -oil immersion.

Capillaria, cgntorta" (Creplin, 1839) Travassos, 1914

pigure-I39, Female. 'Vulva ventral view. -H. p. x IO" Pigure, 140, Female. Vulva lateral view. N. P. X I0. Figure, 14I Egg. -Oil immersion. -, 450

135 13 137

138 i:

3 k 3

140 141 1. oý

139 place the species in his new genus'Pterothominx. Wakelin (1963,: ',

unpubl. ) suggested that C. alpine Bach and Foratner (1959)

is also a, synonym.

First records in British birds are the Black Grouse (Lyrü--

tet ix' (L. )), the-Red-Grouse (Lagöpus' scoticus (Latham)) and the,

Capercaillie (Tetrao'urogallus L. ) from Shipley (1909 a, b).

Subsequently Clapham (1935) found the, nematode in the'Partridge

(Perdix perdix (L'*'))_ and :the Pheasant, (Phasianus colchicue L. ) ), and in 1938she-added the-Redlegged Partridge-(Alectoris rufe L.

the Chukar (Alectoris L. ) the ' Domestic Pigeon''. graeca and .. ,. (Columba liviadom. L. ). Walker (1937) , obtained it from, the _ (Gallus ), ' (1939) Eastern. Domestic Hen gallus L. Baylis, listed'the, - Little Bustard (Otis tetrax'(L. )) and Owen (1951) the Domestic (Meleagris dom. L. ). ' Mettrick Turkey' gallo-pavo redescribed : the species from the (Columba palumbus L. ). The -Woodpigeon, specimens, were obtained from the Woodpigeon, during -writer's ; - . sunnier, spring and, months. In. the Iilackbird-(Turdus merula L. ) only females discovered, (5(1) were -7,11/62). Description

' The length is -between 9 23, (average, gale: :' variable mm. and =. - )I breadth, -13.3 MM- and the is typically 0.036 mm. -across -the -, oesophagus. The ratio anterior to: posterior'part, of'the'body

is therefore 11-1.6. The spicule'length is- also variable, a mm. to 1.682 mm.', and it is slender and ' drawn- to fine-, point, _191 : (Figure 135). For most of its length it is in ý_ _circular. cross-, diameter 0.004 -section with a of mm. to 0.007-, mm. Proximally, `,. -452 it is more triangular broadens 0.009. 0 012 It may and, rto -, . be spatulate or less often irregular, in' outline., The spicule sheath is 1.8 mm. long,, that is larger than the spicule. It, has fine transverse striations and also rows of tiny conical spines.

The caudal end supports a characteristic bursa, 0.05 mm.-in' dianeter, (Figure 136). It, is supported by a'-pair of ¶D-shaped lateral rays, ý0.018 ma. ' long, postero-laterally to the cloaca (Figures 135, & 136). ". There are also a pair of-pre-anal papillae., 0.052 -Anteriorly to the 'bursa are a' pair of lateral -alge, Anm.", long. The shape of, the bursa was constant*throughout a range_ of over fifteen specimens. = in 11mm. Female: The length is even more variable'than the Luale, _ to 43 mm. (average 17 --18 mm. ). The width acrossthe'vulva, is 0.045 am. regardless-, of length., The, vulva divides'the,, body in, the ratio 1-: 2 and is 0.103 mm. from, the oesophagus.,: Most distinctive is-the vulval appendage is tubular and, vase- . which' shaped, 0.068 - 0.085 mm. 'long,, (Figure 137). 'It`leads into a muscular vagina 0.25 mm. long. The tail end is blunt and round. `,, _, ' 0.023, The, eggs are rather, elongated, 0.05 - 0.055, man. by , 0.022 - a mm. 'with fine reticulate outer shell. The inner-shell'is_ reflexed' (Figure 138).

Discussion

No variation in egg length comparable with the, length, of, -the body is Table '33 the ','measurements from :'and spicule seen. --gives , the-descriptions of Madsen'(1945)'and ;i*ettrick (1959)ýfor-, 'longicollis. With the ie'not regard-to the male, spicule. range ,' - ,,, 453. TABLE 32 :C i11ari q cludinfl ata (iiolin, 1858)

Characters Madsen, 1945 Mettriok, 1959 Author

Length 7.0 -20.4 894,11.. 7 9.0 - 23.0 Breadth 0. C25 0.03 0.068 0.036 . Casoph Body'- 1: 1.3 1.1 : - ,-ls1.6 1,682 Spicule length 0.67 - 1.89 0.96 - 1. C' 1.1 - " diaroter 0.005 ... Cross-section - 0 Spiculo tip firm point find point fine point proximal and broad, fraged spatulate n sheath transverse tranzvoroe transveroor striations striations & striations- & fine spines minute apino i & tiro opinesa Burma 2 -- - pro-anal pQp11130-and 2 T_chano(I 0.055 0.095 052 Alae 0. C175- C. (Y77 C.

ona 10 T n3th 9.3 . 36-3 8.9 -! '17.6 11.0- -'43-01, Breadth, 0.03 - 0.07 C. 073 C. C45 , - Vulva rody 1s2 1*2 t - . 0.05 0.1 Vulval ap aondaco more than C. 04 0.6C ... C" 0C; Eggs 0.01x3 - 0.057 0.049 - 0.042 0.05 'ý- 0.055 Z z x 0. CZ2 -, 0.027 0.022 - 0. M4'- 0.022'- 0.023 Outer shell - finely punctate reticulate Inner d- shell reff :. reflexed

455

Anseriformes, Charadriiformes and Passeriformes. = i4adsen,: (1952) ', noted the curious fact that there are no records from gage or domestic birds in'ßritain, and this'was still true in 1962 when

Keymeret al. published their results. There are several records from wild birds. however, Idicfoll - and_ McIntosh (1927) in the,,, . - _. u, .: (Alle-alle (L*). ); Baylis (1928) Carrion Crow Little. -Auk in the (Corvus dorone corone, L. ); Mettrick, (1959)in Crow the Rook -and (Corvus frugilegus L. ) ; Pemberton (196O)In-the Crow land- the , ' ,. Songthrush. (Turdus philomelos clarkii Hartert) and Yemoerton., (1963) in ' the Herring Gull, (Larus argent atus Pont. ), the, Coarnon Gull , . (Larus'canus L. ) and the-Blackheaded Gull(Larus ridibundus L. ). !.

There 'are no Welsh records.

In. this survey specimens have been. recovered from, the. mallard ) - (Aras platyrhynchas L. -ý(3(1) 12/62); the'Curlew, (Numenius arguat t (L" )) - (2(1),. - 12/62); the Fieldfare (Turdus= pilarie L. )

(5(1) --, 3/63); the Robin-(Erithacus rubecula'melophilos'$artert) (2(l) the' Meadow- Pipit (Anthus (L. )),. - -°3/62);. pratensis = (1(1) 'Rock Pipit (Anthus (L. )) (1(1) __7/62);: the spinoletta - , 11/62), (Alauda'arvensis and the Skylark L. ) The,, specimen from the Skylark-was an immature female. ', Description

'mm. gale:,, The length varied between 6 - in the Robin-to 23 " taro. in the Mallard. The average measurement, is. 12.3 an, and acroas the base of the oesophagus is, 0.061mm. No males-were present Pipit,,, the -Rock Pipit,. and'the. in the Meadow Skylark. The '- cuticle fine transverse is thin with striations. The oesophagus varies ,45G from 2 mm. to 7 mm., in, - lengthand divides the body in the_ratioY,

1: 2.5. The tail end is attenuated and bears ,a pair of meagrely developed lateral projections. The spicule could only,,

be satisfactorily observed in a single specimen-where it projected 0.23 mm., beyond the. spicule sheath. The diameter is

0.003 mm. Its proximal termination could not be followed owing

to lack-of chitinization. It is circular ih cross-section.

Conversely the spicule sheath is quite distinct, ' 0.76 mm., to: _ long in Proximally, the, 1.9 " with spines oblique rows. sheath

swells slightly to a diameter-, of 0.027 mm. (for°most ok`, it's

length' it is 0.018 - 0.02 mm. wide). In this region, the ýspines

are uniformly more coarse.

Female: The length'is 14 ma. in the Robin and 30 mm. in 'the :_,. 6 i&allard. The maximum width was, 0.11 mm. -The-vulva is 2: 6.0 mm. - from the anterior end-and 0.084 mm. from the oesophagus. , It'has a, circular aperture which-has a distinctýchitinous'lip,,

(Figures 139. & 140). - The tail is, elongate and slender but ends, (Figure, bluntly. The eggs 141) are 0.054"mm., to 0.059 Anm., by

0.027 mm. to, 0.03, mm. The outer- shell is smooth and the- innere , is shell , not reflexed: Discussion

The spicule approximates to i4adeen's description (1945)

proximal ý eg'e except that, the fraying; , could not be, found. The. all resemble Madsen's Figure 14d, -and are without polar prolong-

ations. ': The Mallard, Curlew and Robin are alle first records in,:

Britain. The Fieldfare (Torgas L. ), Meadow. Pipit= pilaris -the 4 (Anthus pratensis (L. )) and the Skylark (Alauda arvensis, (L. ))-.

are all =new host-ýrecords. Williams (1962) recorded, Capillaries

spp. from the oesophagus of the-Rock Pipit, but this -is_' the. first'

established record for contorts in this host. '

3)" Capillaria obsignata Madsen, 1945-'. `- - _ Syns: Trichocephalus tenuissimus Rudolphi, 1803; ý., , _ Capillaria Trichosomum tenuissimum Diesing, ,ý1851; - dujardini Travassos, ` 1915. __. .. Rudolphi (1819) described T. colombae, from the, large,. intestine

of'Pigeons (Columba, livia dom. L. ) The material' was redescribed.,

by: Duj ardin (1845) as Calodium tenue and particular, reference" was made to the projecting membraneous-a pPendage-rhe. latter was not present in Madsen's material ' (1945) from the"- small `inteat-_ roe, he chose to name species found `in- this locality, and ,-With'o'ut* an appendage, C. dujardini Travassos, - 1915. Travassos -(1914)ý_

', however' had specifically proposed 'duiardini for Calodium tenueý__=

and Read (1949) pointed -out that result duj ardini' as " a .' : .hüst relate to species from the large intestine a vulval, appen ge.. . -with- , of Madsen (1945) differentiated C. -columbae Grayoill`°(1924) - gi8 of shell obsignata on'the basis a reflexed 'collar,, on'the inner , - egg of the and a, differing proximal' end. to the apicule. F, Read=:,,,:" : reassessed, Graybill's', iaaterial and stated : that ,,there' are*-. two capillarids in the pigeon intestine: - large intestine C. (Rud., 1819); syns., Cal. a). - columbae _tenue Dujardin, 1845, Tricho. tenuissimum Diesing,, 1851,

C. dudardini'Travassos, 1914.

b). small intestine C., obsignata ;1adsen, 1945,

In 1951,,; Iadsen acknowledged that Read was correct, and', -. -emphasized that C. '- columbae of general usage is' in fact

C. obsignata. Specimens with a membraneous vulval-appendage--,

e. Trichosoma columbae Rud., 1819, Calodium tenue-Dujardin, ' -(i. 1845, and C. d ardini Travassos, 1914) he synonymised with

C. (Molin, 1858). Skrj et al. (1957) nominated caudinflata , abin, C. obsignata as genotype, but retained C. columbae as, a- distinct

species., A redescription by, udettrick (1959)'was based on Madsen's

original definition of the species 'which was differentiated, 'from, i. columbae-of general usage, e. -without a vulval appendage, and he continued to synonymise dujardini, Travassos, 1915ýwith

columbae on the basis that_the_presence or absence of an_appendage

is unreliable. The most authoritative ; account, - conversant, ',with, ` (1951), dadsen's redefinition - is that of Wakelin (1963).

Owen (1951) records the species inthe-Goose (Anser aneer

dom.. ' L. and, Mettrick' (1959), redescribed the 'species from the

Domestic Pigeon. Pemberton (1960) found the species in the ý. , ýý Feral Pigeon (Columba-livia livia'Gmelin). Keymer et al'. (1962)_ ý_ specimens in, the Wood Pigeon-(Columba L. _.recorded palummbus '), the Stock Dove (Columba L. 'the, Partridge= (Perdix oenas . perdix (L. ))-and 'the Pheasant (Phasianus' ). WakelinIa, colchicus-L. _ -ý- BLANK IN ORIGINAL plate 43

Capillaria obsignata Madsen, 1945

Figure 142 Male. Bursa and Spicule Tip. H. P. x I0 Figure 143 kale. Spicule - proximal expansion. H. P. Z I0 Figure 144 Female. vulva. H. Y. X I0" Figure 145 Egg. tjil immersion.

capillaria ovopunctata (von Linstow, 1873) Figure 146 Male tail. H. P. x I0. Figure 147 Male. Spicule tip. oil immersion. gigure 148 gpicule -.. proximal end. oil immersion. Figure 149 Spicule sheath. H. P. X 15. Figure 150 Female. vulva with appendage. H. P. X I0 Figure 15I EKt, H. P. X 15. , "

a 460

I 148 0 %'

143

...... 146 14/ .i IJ1ci 1

145

4

149 ý 461_ (1963) redescription included, measurement's from the S1ack, Swan )'and, (Cygnus atratus Latham) and the-Hen (Gallus gallus dom. L.

in his thesis-(unpubl.. ) he records the Greenfinch'(Chioris chloria

(L., )). and, the Ashy-headed Goose'(Chloephaga poliocephala Gray). has In. this survey the species ý been found in-, two-new hosts, the ) (8(1) Blackbird (Turdus merula L. - --5/63) and the Starling ) (Sturnus, yulgaris., L. -0(1). -_1/63). Description . .° kale: There', were six males collected measuring 9.5 ---13'. mme' -long The and up to 0.039 u. across the-base of, the, oesophagus. ' divides the body in, the 'of 1 The spicule', oesophagus ratio -°` , long, is is 1.386_mm.. to-1.688. mm. -and-it round'in cross-section,,. -- Proximally it is in-, trumpet-like-`, ', 009 mm. 'wide. - ,; expanded a and curves ventrally, - (Figure ' 143) In this region it `-, , manner .-- . be 0.025'=. in diameter. The distal tip is blunt. The may . sheath, which in four. the 'specimens is everted, `- has ° spicule -of _ fine striations but no spines. ''The-cloaca is surrounded by a bursa supported. rounded cuticular which is laterally` by; a': pair. (Figure 142). There `are lobes of rays, no associated or -alae'.

Female: About twenty specimens were recovered, of which eleven

examined in detail. The length is 17, - 19 mm. --with an,, rwere average widthof_0.055, mm. The vulva, 0.165 mm. behind the has slight lips,, (Figure 144)., It divides the-. body -: oesophagus, : in 1: 2.3. The vagina is rather--short, 0.18- ma. - a ratio ; -long: the caudal end- is, rounded., - The eggs' are 0.05 , mm by 0.025' min.

wide with finely reticulate outer shell. -- The-inner-shell'is 462 not reflexed but forms a distinct collar-round the plugs, (Figure 145). '. Discussion and Table 34 shows kadsen's measurements (1945) those of.

Wakelin (1963). This new"description from two new hosts"falls-, I

the range* of variation, already attributed to the : `ý clearly -within Wakelin's species. The only previous record in a Passerine ho'st'is for the Greenfinch. - This has ' interestsince the Finches are hosts, predominantly seed-eaters., The new Passerine recorded, - herein, the Blackbird and-Starling, are of greater. significance since the Starling particularly is a_winter visitor. Recent 'inease work (Norton, - 1964) has emphasized the considerable in, (up to 49.5 %), in infestation of fowls, with obsignata in the, correlation with utilization of deep-litter ' techniques. poultry industry.. If the Starling is'a natural reservoir host 'infections the Starling 'obsignata :_ - experimental of with are t dealt: with in 'Part 'of this thesis -then the: prevention of, rural access of, these migratory birdsýto the' poultry', houses, in areas is of greater importance than. has hitherto been assumed. BIEý!*s Canllaria obsipnata (Madsen, 1945) . ,. -463

Characters Mad" n, 1945 'Wakelin, 1963 Author

. Length. 6.9 12.96 9.88 - 14.79 9.5 13.0 Bieadth 0.0 2 0.051 0.046 - 0.049 0.039 Ossoph sýBody - - 1sl length Spicule -1.09 1.53 1.43 - 1.78 1.386 ."1.686 diameter n 0.006 0.011 0.004 - 0.009 0.009 Cross-section 0 0 0-_ 'ý - -Spicule tip blunt blunt blunt " proximal end Funnel. -shape Funnel. -shape Funnel-shape sheath tra=verse transverse transverse striations striations striations Bursa lateral rays lateral lateral rays3.: papillae . hength 8.28 - 17.28 13.05 -" 20.5 17.0 ', - 19.0, 0.055 Breadth 0.049 - 0.057 0.053 0.064 .::. -Vulva s Body Vulval appendage none none slight lip Tail end round, obtuse rounded . - -" Eggs 0.04 - 0.056 0.042 0.056 0.05 ' X, 0. a25: ß I I - 0.022 - 0.029 0. ß22- 01029", = Outer shell reticulate reticulate Inner d shell not reflex not retlexed -,. not reflexed 4). Capillaria ovopunc tats (von Linstow, 1873) Travassos-, 1915, , , Syn.: `i'richosoma ovopunctatum, von Linstow, 1873.

This species has been'recorded in the _ßlackbird-_(Turdus meru1a L. ) and-the Starling (Sturnus vulgaris L. ) throughout,: nost ý,.

of the Northern Hemisphere. 'In ' Britain ' it has, been found-in-the . . ettrick'(1959); Starling - Baylis (1928), Evans'(1938'- unpubl. ), ' `and the Blackbird -_Baylis (1939). 'Clapham- (1940) found the

species'in Rook nestlings.. The writer's, material occurred, in

small Walabers in the above hosts.,., Description' .;.

from the Blackbird'_(November, iii: Two 1962), - and`one_from, a Starling'(December,, 1963)., 'The-average lengthýof_these worms: is

0.054 7.3 am. and-"the width' mm. at the, base, of - the ° oesophagus.

The bursa is characteristic 146),. ' with a pair of lateral ': _(Figure ., and, a cuticular dorsal projection. 0.012 'am. in, front ''lobes - are a pair of lateral alae, 0.036 m:n.. -,long and 0.008' nm. wide. The is 0.95, ` -,long tapers-to A, `. spicule mm. and a'-sharp` point. _ spicule nce', hitherto unmentioned' feature of ' the , is the occure of,, -:'-,,

regular annulations ' at` the tip (Figure' 147). ' The proximal

is (Figure 148)-with ' a- slight bias to the ventral -slightly'dilated with side'.,, The spicule, sheath -has marked transverse,, striations ',

distinct ý (Figure -: 0.014 in a pattern ,. 149). It is 0.01-, roan. .`'

diameter. Spines. if present , could -not ' be detected.., Female: Five specimens from' the Blackbird -and- seven: from -the Starling. The body is-whitish-and-up to : 15.7 " mm, long- (from The 'diameter is 0.066 am. " in. the region -Starling). approximately - 46t of the vulva when the uterus is distended with eggs. The vulva ''

is 0.01 mm. 'from-the oesophagus and has an-appendage which may'

be absent or take different forms. Most frequently several, - ý ýit is rather sausage-shaped (Figure 150), but it, may be more squat . or mushroom-like. The largest appendage was 0.13. mm.. 'by 0.059ma.

The eggs have a very, protruding-plug and the lips. are bent later-' , ally. The outer shell has deep striations (Figure 151). and -'.

together with the plugs the eggs measure 0.056 min, by 0.026 mm. ° Discussion

Mettrick's description (1959) is- the ',first - , one .of since. ,. ,-. von Linatow's original. The'details of the male are similar t6,7,

the above text with the, exception that 4ettrick, did not observe the'spicule the annulations at tip. With regard to the female,

the writer is unable to offer, corroborative evidence that , the';, , - -, _, : presence or absence'and shape of_thevulval appendage is, related-,,

to the age. This may be true. The eggs are, indisputably_

striated in - this, collection. In his full text LLettrick describes

them 'as finely 'punctate and in; his key, on-=page '82, he includes with them a raised lattice-like pattern. This contradiction, , -` his -with text ' is' of no significance since - neither' description-; is-', ý- valid. The eggs are, distinctly striated. in' This, is the, first record 'of the worm the Blackbird fron wales. _. -, 5). Capillaria resecta (Dujardin,, 1843) Travassos, 1915 -4661

Syn.: Trichosomum resectumDujardin, '1642; Trichosomä'resects

Baird, 1853; Trichosoma resecta von Linstow; 1877, (b).,

This species has, been recorded from' members of the "family '. Corvidae throughout' the Northern Hemisphere. The British records'' (Corvus are: - the Jackdaw monedula L. ) Baylis., (1928),, Evans (19389 unpuol. ), Trettrick (1959), Pemberton x(1560); the Rook, (Corvus frugilegus-L. ) Baylis (1928), (1959), -Mettrick: "Pemberton (1960); the Jay'(Garrulus glandarius (L. ))-Baylis-(1939); Carrion,

Crow (Corvus corone corone L. 3äylis-(1939); and the Magpie

(Pica pica (L. )): Pemoerton (1960). ` 'The writer, has collected,

specimens extensively from Crows, -Rooks and Jackdaws at -'most - of the-year but especially during sumaer and-early autumn. - -seasons _ Description

male: The description is based on twenty-one readings., The' '10.1 specimens are opaque, - 12.3. mm. 'long and 0.052 mm. 'across the' base of the oesophagus. The is ': caudal region 'rounded withý ". ,ý_ bursa, a cuticular 0.033 -.. 0.036 mm. in diameter, while, laterally_ lobes, 0.012; there are a pair of -the rays of which are mm. _'long (Figure Rook, 152). The spicule is'1.05 --, 1.148 mm. long-in the'' and in. the. Jackdaw ,it reaches 1.265 mm. in length.: The measure- for, the Crows to those the Rook. ', ', It, is . manta approximate of - triangular in 0.007; --' 0.011 but ' ý_ cross-section and mm. across, .; .. distally, it is rounded with a -blunt point. (Figure 152). ' . -ý` - Proximally it expands to"'0.018 when seen "laterally, it , mm., and view, a"spatulate is funnel-like (Figure153). A ventral shows BLANK IN ORIGINAL lamto 44

Ca ipllaria resecta-(Dujardin, 1843)

Pigure; 152 Male tail. Lateral view. H. P. X I0. Figure 153 Spicule, proximal end - lateral view. H. P. X 10" Fieiare 154 Spicule. proximal end - ventral view H. P. x I0" Figure 155 Female. Vulva lateral view. H. P. X 10 . Fibure 156 Egg. oil immersion. ...

Capillaria vanelli (Rudolphi, 1819) ý Figure 15'! Male. Spicule sheath. H. P. x 6. Figure i58 Male. Spicule sheath (a) proximal spines (b) Distal spines 'H. -Figure 159 Female. Vulva appendage 'plaque'. P. X IO Figure 160 Egg. oil immersion. plate 44

Capillaria resecta (Dujardin, 1843)

Figure 152 Male tail. Lateral yie%. H. P. x I0- pi6ure 153 Spicule, proximal end - laterai view. H. P. X 10.

Eibure I'4 Spicule. proximal end - ventral view H. P. x 10. Fisure 155 Female. Vulva lateral view. H. P. x I0- Figure 156 06. oil ima, ersion.

Capillaria vanelli (Rudolphi, 1819)

Figure 151 Male. ppicule sheath. H. p. x 6. Figure 158 Male. Spicule sheath (a) proximal spines (b) Distal spines Figure 159 Female. vulva appenda8e 'plaque,. H. P. X 10 pi6ure 160 E. oil immersion. 4

15i 153

154

155

o\

Iý.. ý. ý

y ý;

.1..

ý/

f. ý!

156 -i _\ 160 158 a. 0-

Q IIIII = 469

shape with a rounded margin (Figure 154).. The sheath is rather,, -',;

longer, than the spicule_and is 0.013, - 0.015 ma. across. It -is smooth along entire length. -. -the Female: Forty-six readings. -The length--varies from 11 mm. 14.25 mm. in the Rooks and Crows and 17.7-mm. in the. Jackdaws., 1

Across the vulva, the diameter, is, 0.073 mm. ' and the oesophagus. is 7.25 5.25 - mm" long. - The-'distance. between=-: oesophagus and -vulva,

isý0.141 mm. The vulva (Figure 155) is, a-transverse, slit, 'without

appendages. The vagina, is muscular and 'up tto' 0.432 mm. ', long. The l is The (57 mm. ýt ai rounded. , .eggs , . measurement s) ' are 0.051 - 0.031 0.058 an. by mm. -with an outer shell which hasýlongitudinal- striations (Figure 156). The inner shell is reflexed'although

this is not always to-be discerned with-ease.

Discussion

(1959)-has, Mettrick described a membraneous appendage'`, in the,

vulvaI region.. This is absent in ' female including all , specimens, - the School (Bottles 3627,, '. 'those'in collection -, Rook: 2695, -; Crow: and Jackdaw: 3704). One must' therefore query-this'aspect

Mettrick's redescription. Otherwise the-, text above agrees well; °. _, with: both, Madsen and Mettrick. Madsen'(1952) considered that , - 1790). not I resecta is, a, synonym of anatis (Schrank, ' ,, This has met- acceptance from with other workers since 'the. egg -shell pattern = _', is quite distinct in'resecta, '(Wakelin'- personal communication). ý-_ In the part ofýthis'thesis. experimental evidence. on-the, inability-to infect-. ducks from will be given. with; resecta -Rooks - . '-., The - species- has not been found in the Rook or Carrion Crow. in Wales. 4-7o 6). Capillaria vanelli (Rud., 1819)'Chabaud, 1952

Syns.: Trichosoma vanelli Rudolphi, 1819; Trichosomum

protractum Diesing, 1851; Trichosoma triloba von Linstow, 1875, ';.

Capillaria vanelli, was formany years-considered-to, be, nomen ý but Chabaud (1952) description of nematode'.. ,, nudum published a a, which is found under the horny lining of the gizzard of the Lapwing (Vanellus vanellus (L. )). He has utilized the name, '

va nelli' for specimens occurring at, this location. ', Von' Linstow - (1875) described'Trichosoma triloba from the same, host and'locatio

This species Chabaud considers to,, be a synonym. - '.. first British-record (1939), The is by Baylis in the Lapwing. - (1959) (as Mettrick published a'redescription triloba) of, this - from the- British Museum together-with somefrom, anew material . -; 'host, 'the Oystercatcher -(Haematopus L. ). The' material-' . , , ostralegus was for this, new description taken 'from =the Lapwing -'(18 1/62,,. (25'(6) '12/62, 12/60, ''1/61, ' -12/62), the Oystercatcher - 7/60, ,, 1/63 (2), 2/63, -7/63), and the Curlew (Numenius arauato$. (L. )) _ .ý (14 (3) - 1/61,11/62,12/62). -, 'All in the Saunderafoo t'' were shot , . - area.

Description

(9 - ', Male: readings). The worms are white and greatly coiled. They only be the. tcan removed with difficulty' since -serpentine .n. S in, convolutions and tendency for `-,the `head or tail-to be'. -buried -:r length',, the. gizzard lining--makes darage very easy. The - average. - m. ' is long is 12.0 n. and width 0.037 mm. The oesophagus, t3.228 - mm. " . ýý " (ratio 1: 3). the of the the ' width ' may -In posterior part . _body''. 47 be - 0.05, m. Striations'-in the cuticle every 0.003 an. occur - The__tail has two lateral lobes, -each with its own papilla, and'.

stretching oetween the lobes'on the dorsal side is a transparent-

membrane. The oursa diameter is 0.028 mm. P-The'sheath is ,.0.85 mm.

long and bears fine spines, (Figure'157). 'Proximally, it is

expanded. slightly and this swelling bears two types of spines., Figure-158 (a)'shows spines .. the strong triangular whichoccupy

, the proximal third ' and, Figure-158 (b) the. finely pointed spi. nels'. 'I

which occupy the-distal two thirds. The; spicule projects, beyond_

this swelling, which terminates in . The ý ý, a, -spine-free membrane. - spicule is 0.37'laan. long and 0.006 -mm, wide,, and 'it is "round in- Proximally the '. is cross-section. . spicule : 'slightly swollen `to -ý diameter, 0.009 'a a of 'mm.. °Distally Ait ýends In fine", 'point. - Female: The females are, 26 31 yn. in length 0.069 mm. wide, -, - and the The is 4.5 ' iving across vulva. - vulva 'mm. from , the front, to-body. lengthof a ratio 1-: 6.88. The distance oetween. theý, = oesophagus and -the is 0.345ýmm.. The isunique in, -vulva vulva - , (Figure that an acorn-shaped, ,' plaque' surrounds ', the aperture `, 159) . -

This cuticular-swelling is seen best in untreated specimens'.

The- vagina is 0: 85 Anm., long. The tail end is rounded, with a.

terminal,: anus. -The eggs-are 0-059, mm. by 0.026 -0.029 '." --'-0-063 . am. and have. a smooth inner is ! outer ' shell, , whilp <

Discussion

Chabaud's measurements ' are' shown with those-of, the writer and dettrick's 'triloba in Noteworthy differences , -Table35. 472 TABIE35* Capillaria vaneelll_i (Rudolphis 1819)

Characters Chabaud, -1952 M®ttrick, 1959 Author,

e Length_ 9.6 10.0 12.0 Breadth 0.045 - 0.037 Spicule length 0-39, 0.72 0.81 0.37 diameter 0.006 0.006 0.006 0.006 Cross-section - .. 0% Spicula tip blunt fine point " proximal and - inflated swollen 0.009 " sheath 0.515 x 0.048 0.43 x 0.15- 0.85 Spination distal 2/3 many spines fine spines fine spines Bursa bibbed bibbed bibbed

Length 31.7 24.0 - 28.0 26.0 Breadth - 0.12 '0.069 Vulva : Body 16.8 1: 5 1 6.88 Vulva - 09s ophagus 0.2 0.2 0-345 Vulval' a PPendage splaquel small circular ýplagweý 0.06 x0. C8 0.05, x0.025 , Vagina 0.9 0.85 Eggs 0.065 0.063: - 0.075 0.061 - 0.057 0.059 - ', x X X 0.025.0.028 - 0.029 ý_ 0.033 - 0.035 '. . o. a26 ': Outer shell smooth smooth smooth Inner shell not reflexed not reflexed not reflexed Tail end rounded rounded , -, rounded -_ý t-z 473

between Chabaud's and this text is the-saaller, spicule length

combined with the greater length of the Epicule sheath. '.. Also,

there are smaller eggs obtaining in the-present account.,, The' , accounted smaller spicule: length may be by Chabaud's statement

that the distal portion of the spicule is unsclerotised. Chabaud

clearly-differentiates, his.. species; from contorta but queries, '',

his male specimen which was not found under the gizzard, , lining but in the oesophagus lumen. The close agreement of'-this text, in from the lining. which all specimens were removed gizzard -, --, indicates that his single male specimen may have penetrated the

horny layer into the The he'male and escaped gut. - characters of'. t of the species is confirmed herein.

Uettrick's description does not show such close agreement.,

He records- a blunt tip to the spicule -and -failed to observe the differences in the spines on the proximal swelling' of 'the sheath. '

4ost'astonishing is-his failure to the'-peculiar mention vulval..: - appendage which he describes as a small. circular,., protruberance. '

This 'plaque' is difficult to determine until the eye is trained, but, can be- demonstrated in untreated specimens. Some, tri lobs specimens from the ßritis-h t useiim, which may, well 'have, been ýý those redescribed. by, Uettrick, were examined and, the, plaque was present.

The, nematode has not been recorded from Wale E; and the

Curlew (Numenius arquatuA (L. )) is 'a 'new' host record. 474

7). Ca, illaria spX"

In the small intestine of a Skylark (Alauda arvensis L. )

-three females of a capillarid species, were' discovered. Two, were - immature, ' 9.5 mm" (oesophagus 4.3 ),, '11.5 mme (oesophagus mn. and . 5.5 mm. ), while the third was mature, 10.5 mm. (oesophagus 5 nm. ). '.

The ratio of the oesophagus to the body-is 1 2.. The body width., 'vulva across the vulva, is 0.056 mm. The, is a transverse slit°

which has a-slight cuticular. swelling- around the aperture,,

(Figure 161). The distance from the oesophagus is 0.028 mm. -'; The. vagina is 1.203 mm. long. The tail end is rounded and the

gnus is terminal. The eggs are'0.052, mm. by 0.029, mm. "'and 'have a striated outer shell.. ' The inner` shell is, not reflexed, (Figure 162). .. Discussion ;-ý..

Capillaria alaudae (Rud., 1819), from the, large intestine of . .' 'the Skylark has not been fully descibed. '' l1ost, characters'

recorded relate to the, male and the ' only details for the, female ' am., are: --length 17 vulva without appendage, eggs 0.052 mm. ' width.,, long and the lateral` bands- occupy. a ,third of the-, body 1 'Although', the mature specimen, is' shorter than the-above-measurement,

the egg length'is is the of a vulval, appen- compatible, as . absence lateral more the dage. - The bands occupy, rather than a, third of-,

width. In the absence species- can, only be, -body -- of a male'-this identified C, (Rud. 1819), 'but'-is the tentatively as alaudae ,- - first British. record, of a capillarid worm in this location in-,, '-",

a Skylark. There are no British, records for alaudae. The bird dissected-in August, 1962'. ' was _ .. BLANK IN ORIGINAL -daue 45

Capillaria sPP" (Skylark)

Figure 161 Female. Vulva lateral view. H. P. 'x I0" Figure 162 Egg. Oil immersion.

Capillaria spp. (Twite)

Figure 163 male. Bursa and distal end of spicule. H. P. XI0"- rigure 164 male. gpicule. proximal end. H. P. 1 I0"- 476

4 78-, Subclass: Secernentia (von Linstow, "1905) Chitwood, '..1940

Syn.: Subclass Phasmidia Chitwood & Chitwood, '1933 Class, Secernentea (von Linstow,, 1905) `Dougherty. ',,1958 , Order: STRONGYLIDA

Super-family: STRONGYLOIDEA (Weinland, 1858) -Hall, 1916 _ Family: Syngamidae Leiper, 1912,

Genus : Syngamus Siebold, -1836

Syn.: Cyathostoma Blanchard, 1849

9). Syngamus trachea (itontagu, 1811) Leiper, 1912-

Syns.: Fasciola trachea Montagu, '1811;, Syngamus trachealis Siebold, 1836; Strong yluspictus Creplin,. 1849; yngamus bifurcatus Theobold, ` 1896;. Syngamus mucronatus

Schlotthauoer, 1860; Syngamus primitivus' Molin, 1861; Sn amus . sclerostum, Tdolin, ý 1861.

Half dozen pairs in copulq were recovered. from the trachea' -. ,a , . of-a young Rook'(Corvus frugile'Kus L. ) in July, '1962. This species has-had numerous records from Wales both, in, poultry-_and'-: in wild birds. It would appear to: be, unimportant. in Pembrokeshire . however, from ' the migrant birds: and-it was not, recovered any of . such as the flocks of starlings, in winter, which might be to be introducing_a parasite of economic importance to'--, -,- 'thought local stock. - The species is too, well-known-, to, warrant yet another redescription. ' Order'ASCARIDIDA

Supelr-family: Ascaridoidea

Family: Heterocheilidae Railliet et Henry, 1912. " Subfamily: Filocapsulariinae Yamaguti, 1961

Yannaguti (1961) nominated the subfamily Filocapsulariinae

in place of, Anisakinae Railliet et Henry, 1912, because Anisaki'

has been proven a synonym of Filocapsularia Deslongchamps,,, 1624. -,

Genus: Contracaecum Railliet et Henry, 1912..: _ Syns. -: Kathleena Leiper and Atkinson, 1914; 'Cerascaris'Cobb, 192

collect- The two genera from the, Ascaridoidea in, thepresent

ion are easily distinguished by the presence= of an 'oesophageal (1959) appendix. ' Osche has, given a 'comparative account, of the

morphogenesis of the lips of Contracaecum and Yorrocaecum. `

10). '-'Contracaecum spiculigerun_(Rudolphi,, 1809)__' Railliet Henry, 1912' et . Syn.: Ascaris spiculigerum Rudoiphi9', 1809, '- _-ý

British records for this universally dispersed species-

are:-the Cormorant (Yhalacrocorax carbo (L. ))- Lewis, 192?;. Baylis, _ (unpubi. ); '1961: 1928; Davies, 1937 Baylis, -1939; -Williams,

the Shagý(Phalacrocorax ariatotel is, "(L. )) - McIntosh & Nicoll', -,'- 1927; Baylis, 1928;: 1939; 1961';, the, Green Cormorant,. =( -Williams, ) -Davies, 1937: "(unpubl. ):, 'the , Black-headed 'Gull';. & (Larus ridibundus L. ')', -, McIntosh, & Nicoll, 1927;. Jennings `.., ' -. 1958: the ' Guillemot (Uris' (Pont. )) dcIntosh '& Soulsby, -- aalg, te -, Nicoll, 1927.. The, ' recovered they _.ý writer's specimens ,were -from; . -: 480 Shag - (8 (1 ); -ý 11/63) ; the Razorbill" (Alca ' torda L. )

(1 (1) - 7/62); the, Guillemot 7/62); the Lesser_, ': Black-backed Gull (Larus fuscus-L. ), (2, (1) 7/62) and- the, -" ,- --,,,-. Manx Shearwater (Puffinus puffinus (Brlinnich))- (1; (1) 7/62). ' There three in the Shag in the Guillemot.. ' were males . ,,and one' The remaining female both those from the Aanx -, specimens - are . _'arid' , I'.I-'- Shearwater. and Razoroill were immature. Measurements" derived

from various-sources are stated in brackets after the writer's

own. -

Description - {lam: Male: The 19.6 and 24.5 long, ' specimens were mm. , -23-0 mm. ", '-, The average breadth= was 0.85 mm. "_(20.0,, - _45.0 by 0.014- 1.001 min. ). The head is quite distinctive, (Figure; "165); `equal three large lips with` conspicuous of nearly _interlabia length-and-an overall diameter of 0.24 min. are followed by. ýe , folded deeply cervical ýe, xpansion, " 0.289 =mm. " across. - "Iauediately, . , ., posterior to "this "a' deep transverse furrow, ' after, which 'the _is ,_°° body thickens to 0.365 an. The cuticle " is'. 0.019 thick and', "ý

the 0.005 min. " The lips striations" are . , apart. are-0.115", mm. across and by ma. 0412 m:n., long (0.075 -0.139 0.085 0.117- ) rend _. - _mrn. " the 'papillae are 0.013 in diameter. -, conspicuous, mm. " "A= dentig- blunt erous ridge is absent but, anteriorly from each lip _,' horn' _ . , .a -protrudes for 0.026 mm.. The interlabia are 0.082 min. " long,;, ' (0.061 - 0.112 by 0.036 0. '074. mm. ) ýnerve ring is _mm. -. . _'--The"" . L- 0.458 mm. from' the ° anterior end (0.52 -' 0.65u mm. ). The

oesophagus-is 2.24 long (3.0""- 5.2- mm. ) ' and the width ,'-3.06 mm.°: ,. . .ýc 481 is 'approximately 00.17211'='; The ventriculus is not very distinct, ý248 226 (0.226 mm. ) 0. ,qmm. 'byýi, O. °mm: - 0.375 by 0'. 2, -'0.32 MAU. ý_ and Xrom''it arisesla ventricular appendix, 0.89`mm. =by 0.152mm. (0.714 - 1.3-ma. by' 0.096 - 0.25 mm. ). ' The intestinal caecum mm. Js 2.24 by-0.21 mm. (2.03- 3.3, mm.' by-0.192 am. ). - The tail (Figure"166), is: conical and ventrally curved'with` 'I cloaca the 0.212° mm. from the posterior ýtip. The -spicules are long, half 'the (Figure " extremely 'almost body length -167), equal - and slate. The exact length of those figured is -11.763 ). (6.2 - 9.1 mm. The shaft is 0.019 mm.:, across and with the alae' thie total width is 0.047, am. The end is pointed. d. -, The ' number of pre-anal papillae is difficult to estimate, but there fewer than, thirty are not pairs. ýThee seven post-anal pairs _ _are arranged'as_, shown in Figure 166, 'namely three lateral 'pairs, of.,, which the middle pair, is the smallest. ` and; four subventral r pairs, two of which are, closer to, the cloaca, '. -pair 'median - -'with one -to; ' ' the two the `-`. the others, and rear pairs adjacent to. -first' and::, ' lateral second pairs. . Femme: (Five- readings). The body-is-52.0 mm.''long and 1.05 as, wide (36.0 -85-0 mm. by, 0.96 1.8"Mm. ) and the, head, shows

similar organization to that of. the male. ' The lips, 0.119 mm: -'are ,. _:; (0.11 by 0.11 mm. - 0.151 by 0.11 - 0.131 mm. ), and the

interl ab is are 0.085 ' 1=. by 0.07 Mm. (0.111 ` mm. by 0.072a, n. The head diameter, is 0.259 min. (0.28 - 0.42wmm. ) and it foil- 'is owed by, a striated cuticular, expansion, 0.328-mm. across. ':, " The'-' nerve ring is 0.63 mm., from the anterior (0.62 0: 76 end . 482

iIiV E="! d31ý1 , 0; '''ll 0'!. The ie, a3. 751 by 0.28 (3.5 by 'pi ! oesophagus' li y mm. mm. ' 5.3 nm. 0.2 ýý ýIt - . ý'and ,j0.31 mm.') leaäs'ý into' a ventriculus 0- 32, by 0.318 am., (Figure ' 168). The ventricular appendix is 1.128 0.041 . mm. ,b. am. (0.84 0.14 0.21 ) -with -X1.5 am. -by - m,n. , an intestinal caecumr. (2.4, 2.872 mm., by 0.314-mm. - 4.4, mm. by 0.203 mm. The vulva '

is in the anterior third of-the body and divides-the'bödy,, in the. 'tail ratio of 1i2.6. The tip is conical. and, the anus is

0.284 mm. from the' posterior, end.., ' The lateral papillae-mention'ed'

by Yamaguti (1941) could not be discerned. The eggs-are sub-

0.072 mm.' long end. 0.061, is spherical, mm. across. _The, shell',, 0.004 mm. thick and the ovum unsegmented. ', Discussion .. _ ,: The specimens fall well within the range. ' of recorded'--

measurements. The only significantdifferencJes are the extreme

length' of the spicules in the male, almost half that " of -the body,.

and the apparent absence of the 'lateral papillae inthe female There has been tail. : no redescription from material 'found-'in

Britain. The'Razorbill, Lesser'Black-backed Gull and Manx' British ) Shearwater are-all new ' host , records.

The life-cycle was described in part, by Thomas '(1937). BLANK IN ORIGINAL plate 46

Contracaecu, a spiculigerum (Rudolphi, 1809)

Figure 165 Dorsal lip. H. P. X 10* Figure 166 Male tail. H. P. X I0- Figure 167 Male tail with spiculea. L. P. X6 Figure 168 gemele. Oesophagus. L. P. x 6.

-porrocaecum crassum (Deslonchamps, 1824)

-Figure 169 Dorsal 'lip. H. P. X JO. Figure-770 Heau end. L. P. x 6. 484, Genus: Porrocaecum Railliet et. Henry, 1912

Syn.: Terranova Leiper and Atkinson, 1914'

The genus is well established lin fishes, reptiles, birds, , , marine mammals and insectivores. -Apart from the-absence, of an

intestinal, caecum-the presence of dentigerous'ridges is, charact-

eristic. The-lips-have diagnostic value at a specific'level. ý.

also. The dorsal lip has. a pair of sub-dorsal double papillae'-",

while the lateral lips each have a fully-developed sub-ventral, -.

papilla and a vestigeal sub-lateral, papilla.:. The interlabia=;. ',

rarely more than extensions the, but the lips are . of . cuticle body tissues as lobes contain a' pulp' ... Anteriorly; a l,pair of arise which are variously subdivided into -anterior, and lateral'

lobulii, '(Figures 169,171,176,181). ' On the inner°side-of lip is lobus"impar, the each the -`-sometimes- projecting beyond, ' ' lip border and sometimes not . The species from European birds have been-studied and ... key provided by Hartwich' ('1959);

L1). Porrocaecum - crassum (Deslongchemps, '- - -1824) . ._ Henry, 1912: Raillietlýet _ 1824 Syn.: Ascaris crassum Deslongchamps,

The type host is the, domestic duck (Anas latyrhynchns dom. °. same: L. ) and the first -British record occurred in this host in Lewis (1926). Baylis (1928) ` the species the i allard --- recorded ,,. (Aras I;. ). Subsequent fron the Domestic platyrhynchas records °:- Duck are from Wales -'Davies (1937 - unpublished) and. Evans ',

(1938, - unpublished). ', Two immature females were taken from the Mallard (1) - ý.:. "_ -(2 -12/62). Description

The measurements recorded by, previous authors -_DuJardin

1845; Skj abin, : 192 j; ' Mosgovoy, 1953 and Hartwich, 1959 "-- are:

stated in brackets , following those. of the writer. The average -', .; length these is 18, ) the of specimens mai. "(43.0, -'66.0 mm. and (2.0 2.2, ). E breadth 0.58 mm. -, - mm. , 'The head diameter is 0.274, m. - (0.334 mm. ) and the lips '. dentigerous --0.584 are conspicuous with- ., ridges which extend to the-respective sub-dorsal ýor sub-ventral'. _

papillae_(Figure, 169). The lobes of labial pulp are broad-'-:, -`, _the and at the inner edge are three short digitate-projections.;

Laterally the lobe has, a small extension, - the-stall piece'. , Figure, 169 ' shows the dorsal lip' the and, part of - , right_, sub-ventral lip. The median lobus impar projects only a--very'short distance,, and, beyond the main, labial pulp. -The;. interlabia are small` , .,: pyramidal, approximately-two-fifths-of, the lip ! length. Cervical, - . ý, alae are absent. . The, oesophagus (Figure 170) is 1.21' inme long ,-4.65 mm: , -(3.4 and. leads" to by 0.254 . `. ' a_small oblong ventriculus, 0.212°mm:. . mom.-ýý (0.4 -=0.462 mm. by 0.653, mm. ), The intestinal caecum, is 0.69 mm., long (1.414 '-' 1.9 mm. by 0.3 54 ), that i8 three, to times- -, - inn. . - -six the length of ' the ventriculus. ' The vulva is In the second third

the body. 'ý The is the is 0.226, =o'' from,,,,, of

the posterior tip. There were no, edge. 487 . Discussion'

The short ventriculue, -broader than-long, -is characteristic of this species, the genotype. Similarly the length of the, intestinal is three to five times that of-the ventriculua _caecum ll (1959). Thus and the lips correspond to Hartwich's redescription _, despite the small and-immature 'specimens, 'it is possible to

provide-an accurate-identification.. ' The-lateral papillae'-which-ý:. Hartwich describes between'the third-and, last quarter of'the tail,

discovered. The Mallard is a host record, Wales': were not . -new -for and is the first from -a non-domesticated "This is also the' , -bird. . from'British ' The life-cycle first redescription material., . was studied by Mosgovoy in 1952. _,.

12). Porrocaecum-ensicaudatum (Zeder, Baylis, 1920 -1800), Syns.: Fusaria ensicaudatum. Zeder, ' 1800; Ascaris . '1809; ensicaudatum (Zeder, =1800) , Rudolphi, , Asceris- crenata ,, (Zeder, 1800), Rudolphi, ': 1809; Fusaria' lancea Zeder, ; 1800; °" -Rudolphs, Ascaris lancea (Zeder, 1800), 1809; turdi; sSpiroPtera, Molin, 1859; Filaria Linstow, 1877. ' - turdi 'von -. '. and The type host is the Blackbird (Turdus merula L. ) Lewis, British records from'-this host are: --Bellingham, 1844;. 1926,, 1927; Baylis, 1928; Davies, -'-(1937 unpubl. ) ; Baylis, 1939: -- and- Mettrick, 1960. _, Ther first British record occurred in, the

Missel Thrush (Turdus viscivorue Lo )q Bellingham., 544I ° Baylis, 1928; Evans-(1938 - unpubl. ); and Jettrick, 1960. Other members of the, family Turdidae to be hosts. in Britain are the Songthrush'(Turdus L. ) Baylis, 1928;. Baylis,, . philomelos - . -` 1939; Jettrick, 1960; riilliams,. 1961:, 'the Fieldfare (Turdus

pilaris L. ) - Mettrick, '1960: the" Redwing (Turdus musicus L. ) - -Baird, 1853; Baylis, ' 1928 1939: the, Icelandic Redwing" and -and (Turdus musicus 'coburni ?)- Fcott, 1957". -- .'; There are numerous records for Corvid birds - the Rook,. -There _ (Corvus ) frugilegus', L. -Lewis, '1927; Baylis, 1928; Evans-(1938 - ); and , unpubl. Baylis, 1939; Mettrick, -19604 Pemberton,, -,1960 Williams, ', 1961: 'the Carrion Crow (Corvus corone corone. L. )

Baylis, 1939 and iAettrick, 1960; ' the °Jackdaw' (Corvus Laonedula' L. )

- Mettrick, '1960 and Pemberton, "1960: and the Jay (Garrulus', (L))' glandarius - Mettrick, 1560. ' Records. from the: Starling (Sturnus vulgaris L. ) Lewis, 1926,1927; 1928; Walker, -' Baylis, . - 1937; ' Evans, (1938 unpubl: ) ; Baylis, 1939 and '. Aettrick, 1960-' _- - . The writer's material was found in-the Rook--'(25 (11)--: 19609 61,62,63),, the Carrion Crow (5 (1)-'1/63), the Starling (8 (2) -12/62,1/63), the Blackbird (9 (3) - 4/61, ' 1/62,, =5/63), ' the, Redwing (3 (1) 3/63), the Fieldfare (5, (2), -, the - -'-3/63),. - (6 (1) Songthrush -'8/63) and the Black-headedýGull, (Larus ridi*oundus L. ) (2ý(1) 1/63). All,. in', - - -, were examined' ° Pembrokeshire.

Description (Nine and Male: readings). The length- is -28.0'- 35.0 mina the

breadth is 0.58 - 0.73-mm. The head. diameter-*is 0.47 mm;, BLANK IN ORIGINAL plate 47

porrocaecum ensicaudatum -. (Zeder, 1600)

Figure 17j Dorsal lip. H. P. x 10. Figure 172 Head. Anterior view. H. P. x 10" Figure 173 Intestinal caecum. Larus ridibundus H. P. x 6. Figure 174 Intestinal caecum. Turdus pilaris L. P. x 10" Fi6ure 177 Male tail. ri. P. x 10.

}orrocaecum semiteres (Zeder, 1800)

Figure 176 Dorsal lip. H. P. X I0" Figure 177 Intestinal caecum. L. P. X I0" iie; ure 178 male tail. L. P. x I0 " Figure 179 SPicule. H. P. x 10" 490

176

m o. ýmm

179 175

o

0

0

0

0

0

0

0

E E "O / 05mm ..

/O /O000 0r while those' of the ventriculus and tail are 0.616 and 0.21 m.a. . There are three 'lips, oval ' in, outline and'with a'distinct-cleft or the anterior edge of the dorsal in 'specimen (FFigure ' 171). - , . each The_ length is 0.132 mm. and the central pulp is oblong and solid''

in, outline. ' The lobus, impar on the-inner side of each lip does'°:

not project beyond the central pulp but may be seen in the.

anterior view (Figure 172). Each lip bears a pair of frontal

lobes which are " distinctive, to this species and"-the closely, ' Anteriorly related semiteres. there. is'a, -cleft between two lob-' ules, 'while a distinct lateral lobule 'tailpiece' extends . as a `to the papillae (Figure. 171). does almost The' dentigerous , ridge ., not extend to the papillae. ' The interlabia-'are conical'--, expansions

of the, cuticle, half to *three-fifths, 'of the lip, length. - There ý. , . are no cervical alae.

The-cuticle varies in thickness from 0.009 015'awa. - with -'0 . ._ 0.006 is striations -0.008 mm. apart. , The ' oesophagus length by 2.03 -, 2.18mm. while'the ventriculus is 0.428_- 0.49 am.

0.132 - 0.284 an. That of the' male in, the Black-headed"Gull'

measured 0.175 mm. by 0.132-mm. 'The-intestinal-caecum in-the ''-

nale is never more than third the length the ' one of . ventriculus`'. (Figure"174) except in the black-headed-Gull'. where'itis', almost`, -

as long, 0.169 am. (Figure 173). 'The'tail narrows, to a_-small (Figure not',. cone 175) with'a thick, cuticle which does however, _,

expand to form a caudal alae, proper.. The cloaca, is O'27-mm.

from the posterior tip. There, are 14-18 pairs of, pre-anal apapillae

" and six pairs of post-analL, the'-first of : which is a subventral'., 491, .. double-pair adjacent ' to the cloaca and-0.249 min. from the 'posterior . end., A short gap separates the second, and fourth pairs -third which are eonverginß 'towards ' the' axis. -subventral midventral, ' -ý , The fifth -pair are lateral. and the sixth are 'again subventral,

0.072 mm. from the posterior' tip. ' The spicules' are'slate, 0.55-

'0.69 mm., long. In the Black-headed gull: they, are only 0.48 mm. (Twelve Female: readings). The longest specimen-was 43-0 breadth 1.55 The with a of mm. average diameter of-the-head is-, - --, 0.323 mm.', across the ventriculus 1.3 ax., and of. tail nn. _the _0.35. The cuticle is very thick, -, up to 0.31 mm. in the ventricular °.". . - '_ The lips male, region. are similar to those of the -0'0212'=. long, 'and the interlabia 0.119 'ong. is are yam. , The oesophagus, 2.54 -ma. long and ' the ventriculus is 0.49 m,a. -by' 0.297. mm."' The intestinal caecum is-0-332 ma. -' long and never exceeds the length ,,., of=the ventriculus.. 'In the longest the-vulva is specimen, _<< 22.0 ý- am. from the anterior, end, i. e. -Just within the anterior', half of the worm. The tail is blunt 'and' the anus 0.63, lam. from, the posterior end. In the terminal fifth are a pair of,. lateral .-'. papillae. ', The eggs have an outer. shell with a reticulate pattern,, 0.099 am. by 0.063 am.

Discussion"

The specimens from 'the Songthrush iLalnature, found were all', .; the horny ' layer. to' Alolin under of the gizzard. - They correspond Spiroptera `turdi (1856), which Mawson (1956) showed to,, be the , 4th.: larval stage 'of ensicaudatUM. This has been confirmed by '- , Levin (1957) and Osche (1957).. Table, 36 gives the, author'a CO D r' 14 Cp% O O r N N M M C4 M O cil " " " " " 1t1 Oý q- , O, O O M M M O, nN ' O IýO O O O M O N O Lr% .ý tit co 11 0 Ö C ö d öcý ö CN N i 1 !! ý. ýC 11 " ,-+ 0 : c3 0 1 1 1 1 _1 O O O CC) _ ý O ý CO r " %o Cy cr% 0 O 0 O N .4 0 ' O_ 0 cV 0 O O gyp 0 O _, measurements alongside those ofHartwich (1959) and Mettrick,.

(1960). Every measurement falls within those'already recorded. for the species with, the 'single' exception of the intestinal caecum in the male specimens from the Black-headed Gull (Larus ); ridibundus L. these are unique" in being , almost the length :of (Figure 173). Osche (1959)-on, describes the_ventriculus -page 468 Porrocaecum larvae 'in 'Fehlwirten' = false hosts- -and' in-, particular the Black-headed, "Commonand Herring Gulls, and'the- woodcock-(Scolopax, rusticola L. "). From under the, horny, layer - of the, gizzard he ' has described , larvae ('Infective larvae') in"- - , the Black-headed Gull, which he'is able to compare ' directly with". `', intermediate-host, -',the; ensicaudatum'from: -the correct earthworm (Lumbricus terrestris). from the-adult --This-third etagediffers in lip morphology. and other features 'so ' different host`--, and -in .a , and a ,.vertebrate' one also,. there is. no reason' to suppose, that-_ further development would occur.,, ', However in the Herring- Gull,

Osche records ensicaudatum larva -1V of `,which seven 'out - of °' _ -twelve were living., The only previous record: which indicates 'full " .. ý_ development in, the family Laridae, is from. Pemberton, (1963) -. who ,. found Porrocaecum species, in the Herring Gull and the Black= records. headed Gull as new host, The ' Black-headed--Gull as 'a`: "ý" definitive. host for ensicaudatum is confirmed in ` this', text ae, _; a new host record 'in so far as '.the -specimens can-be identified. ---_, The small size'of the spicules. is, more suggestive of ensicaudatum' than of semiteres although-Lewis '(1926) has queried the, value .;; of spicule length as, a. criterion in distinguishing-between these. A9 two species. -Despite the length of the intestinal caecuin, because it does not extend beyond the ventriculus thespecies is more obviously ensicaudetum than semiteres. The writer's-"; ' main identification former species is- based with-the . on ecology rather than P. semiteres 'is ,found in the family.; morphology. --'. ' . Charadriidae, are insect-eaters rather ° than ; which - eaters of earthworms. (Based the own findings in. the . -on -writer's -gizzards. this and crops of birds dissected during survey-, -. careful'_

correlation of_parasite and gizzard content always being made. )

The commoner Lari dae are increasingly 'land-bound I and ' during the

winter 'months eat enormous, quantities of earthworms, and a', feIature--> of commented on more fully in the-. analysis this survey, is,,., -to -be, ýin the loss parasites the winter; 'months-, when` general , of usual fully, to _tIerrestr- the diet changes from being littoral or marine

ial comp-,onents. However a nematode species which'usesthe'earth- itself in worm'as an-intermediate host-_couldeasily establish: a.. of non-passerine host during the temporary change diet. -- Osche the larva. of seraiteres -(1959), p. 466-in discussing -stages -has larvae in the intestine found larva III with 'ensicaudatum of the Field. ma=als the Water Vole (Neomus fodiens) and Mouse (Apdemus in-his inference that agrariu). He-is mistaken hosts must have obtained;; these hosts and the definitive'Charadrid (top, page 461) since both the 'semiteres from Oligochaetes of , insectivores (eating mice and the birds are more specifically. stages) an insect intermediate-host both larval and adult and is more certain. Hartwich (1959) raised a new combination P. picae (Rud., =

1819) for Porrocaecum app. from, corvid birds., His main

criterion, is the intestinal caecum,, which is up to, but never, ventriculus. more than, I-one and a half times the length'of the

He-places the ensicaudatum of. Baylis_(1939) under. this species. iVhile' time to Baylis"material, ' there has not-been examine none . collected=from corvid birds during this work possesses so long..: feature found, the a caecum and until -this particular -is writer, is exercising caution in. accepting the new combination.,

The Gull' is a new' host record for the species,,, _Black-headed' while the Carrion Crow, the, Fieldfare the Redwing are ° all -and` , ,, first records in Wales. -

13). Porrocaecum semiteres1800)'(Zeder,, Baylis, 1920

Syns.: Fusaria semiteres Zeder, 1800; --Ascaris semiteres (Zeder, Ascaris, 1829; 1800), Rudolphi, 1802; , heteroura Creplin, " Porrocaecum heteroura °(Creplin, 1829), 'Baylis,, 1920, Asp

retusa Fröhlich, 1802.

The Lapwing (Vainellus vanellus L. ) is the type host. and

the species has been, recorded by Baird_, (1853),. McIntosh-and. Nicoll (1927), Baylis-(1928), Davies (1937,, unp ), and': Nico11-_(1927) (1938, unpub1. ) and -Baylis'(1939)... -, McIntosh' Plover. -recorded-Ascaris heteroura Creplin,, 1829, 'from-the`Golden (Charedrius L. ). apricarius _ _ The writer's in, the' Lapwing_: '- material was recovered ,, 1/62) Plover (1 (1) and the Golden - -, 1/63).: The Specimen from the Golden Plover was a male. The measurements ,-ý Of Hartwich, (1959), follow those of the , writer. Description

dale Two readings., ' The male, from Lapwing wash damaged, ao ' _the . .` the following measurements' relate to the male' from , the 'Golden . , , , Plover.. length 24.0 the breadth is. 'O. 8.54,, The is '_mm. -and - 0.934 ,, , . ). LQ4" (22.0, 140.0 by 0.56 1.0 MM. The- head, mm. - , min. - -is-'0.235 ,ý .- across, the 0.66 mu: and the tail, 1.15 ' mm. The ventriculus _ m. cuticle is 0.015 mm. wide and has striations 0.006 n. apart in. body. The the neck region and 0.012 mm. on the cervical- al as, ':

recorded by several workers were not discerned.

The lips are very-, similar to those of ensicaudatum but are- than and wider than long, ', 0.149 mm,`., by -oblong rather oval 0.198 The, edge of the dorsaL. mm. (Figure 176). anterior ýlip ,ý lobuss is the described, in ensicaudatum and the imp ar -without cleft pulp'of the 'lip, ie does project slightly beyond the main _`which- The llob_es=andjobult 'are rather' narrower than in ensicaudatum. The interlabia are rather more identical with ensicaudatum. - , least two-thirds of the lip length. strongly developed, - at aim and' The long` (1.76 .) ,the;.,; oesophagus is 1.63 man* ,-2.05 in "- 0.346 ventriculus less elongated than, that - -ensicaudatum, The, intestinal caecum a : never by 0.306 mn.. (0.22 - 0.375 ý") " ý_ the length of the. ve ntriculue ý_ , less than one and a halftimes . . , 49'

(Figure 1?? ) be double this iength, '0.476'mm. and may -being recorded for this material (0.24 0.65 ' mm.'). - The is - . cloaca" 0.33 mm. from the posterior= end and the tail is 'conical (Figure'::; ." . 178). Thirteen pairs of pre-anal-papillae were counted and'six' pairs'of post-anal. These show-virtually-'the same distribution, -.., as those of ensicaudatum, with tha first-pair'immediately`poet- '

double', the fifth laterally The- ';. anal and and pair placed., - spicules are alate in the Golden Plover, 0.645 ma. long-andin' -a the Lapwing 0.86 mm., long (Figure 1? 9), (0.605-- 1.24 mm. ). =

Female: The four 32.5 38.0 47.0 specimens' were -' nm. , ,mm. , mm. (21,0 ). and 55.3, mm. respectively, -_58.0 mm. The breadth :. =. _long is 0.658 M.U. (0.625 1.45 am. ), The oesophagus was 1.862 iau: - , , by 0.298 m=a. (1_. 5 2.98 ) the 0.27 0.312mm. -. - mm. and ventriculus. - 0.346 by 09332- mm. - wider, than long, unlike the , males. -,The caecum. is"0.332 mm. to,, mm.. long (0.221- 0.62 mm. ). The -0.525 vulva occurs 'between-the second and-third fifths of the - body ;Fand , - the sculptured reticulated eggs , are 0.085---- 0.109 min. ' by 0.063=, -, 0.066 min. (0.093 'mm. by 0.07 ). There ' is lateral' , min. . °a' pair, of f-- alae Lnear the tail tip. '. ' Discussion (1922)- Baylis separated-semiteree from ensicaudatum. His-, -, criteria, despite -criticism from Lewis (1927), remain valid,: ae..: is demonstratedby'a comparison of the above descriptions with to, lips and, caecal length. '- Lewis (1927) regard r-recorded semiteres in'-Rooks-and Starlings. If the. lengths Lewis' material ` caecal . of is closer, to'semiterea-than to, ensicaudatum, then, it. is-possible' "A 99 that Lewis was describing the species which Hartwich (1959)' believes to- be, P. ipcae (Ruda, 1819). Baylis stated. that., thet . so two species are really little more than subspecies. Even the distinctive features of_each remains remarkably constant through- out the range if. the various European, ' American and Russian

are redescriptione, -valid. ,-. The Golden Plover is a new host for Wales.

14). Porrocaecua (Rud., 1795) Baylis. '1920 spirale . Syns.: spirale Rudolphi,, 1795; Ascaris depressum, 11 -Ascaris t'= (Zeder, 1800) Rudolphs, 1819; Ascaris bulomus Prbhlich,: 1802. ý

The original. material was found by Rudolphi in, the Tawny ) Owl (Strix aluco L. and , this ý host was recorded' by_ýi3aylis, w 1939-:: ",;

Other British hosts are the Long-eared Owl (Asio otus (L. ))--_

Bellingham, the Short-eared Owl (Asio flamme `us (Pont. ) - -1844; _ ,:, the Barn, Owl (Tyto, (Scopolli)) Baird, Lewis, --1926; albs' '. 1853;

Baylis, and the Little Owl noctua mira Witherby) -' . _1939; _(Athene ' 1927; Baylis, 1928; and Baylis, '1939" The-mate for -Lewis, -. ri al this redescription was found in the ' Little Owl C3 , and the Barn Owl -(1 (1) - 11/63). _ ,- Description, '

Male: One reading. The is 39 0, mm. and the breadth, "is _,_ -length 1.1 The, head diameter is 0.398 mm:, while the ventriculus '., . BLANK IN ORIGINAL a plate 148

porrocaecum spirale kRudolphi, i795).

Figure 180 Anterior end with caecum. L. P. xb Figure 181 Dorsal lip. H. P. xb Figure 182 Male tail. L. P. X I0"

gtreptocara tridentata (von Linstow, 18'13)

Figure 183 Female. Anterior end. H. P. x I0" ifigure 184 Egg. oil immersion. 5e1

»

181 184 I nß r'' ýý I° E

O 182ýý -ý'

E E

0 and tail-are 0.714' mm. and 0.24 min. '. across respectively. The >> , lips hexagonal 0.264 long. The pulp=is with are and am. , oval two anterior lobes, cleft into-two short-lobull'(Figure 181)., '

The lateral lobule-or. tail-piece is much reduced, =not reaching the The lobus impar is projecting to papillae. - well-developed, for at least, half. the length of the anterior lobes.,, The -,dent- and igerous ridge-extends'to-near the stem of the, lip, the conical-'- interlabia 0.098 long. ý than third of ý the are am. - rather more .-a' ' -; lobe length. There''are- no cervical'alae. ° The cuticle is', 0.027mm. thick with striatlons up to, 0.012 an. apart' in mid-body. The and 'oesophagus is 1.6 mm'. long the ventriculus oblong, 0.314 .nm., oy 0.346 mm. The intestinal caecum, is three times as long, as

the ventriculus,, 1.08 mm. by 0.153 mm. (Figure 180). The, anus _,, 0.232 from -_, is mm. the posterior, end. The pre-anal, papillae number about pairs.,, " There gare six pairs of , post-anal twelve papillae of -which the first. pair is doubled, (Figure 182). The remaining five 'pairs are I all on. the small conical tail-piece, ',. ''

the first and fourth being subventral. The -:third and ,,fifth 'pairs

are ventral, and the second pair are lateral., They,, were not ý, , -=conspicuously asymmetrical ' as, indicated' by'Hartwich (1959)"

The spicules are elate, 0.58 -, 0.64 mm. long..

Female:, (Three readings. ) The -length is59.0 mm.. and the,:. =ý:_.. "

breadth-1.4 am.. Most other measurements'are. -slightly' larger, than those of they male. The is ''in the posterior half. , vulva vagina It has distinct transverse lips and the muscular, is -=ýý 0.43 mm. long. The anus As 0.414° ma. ', from the posterior tip

Tait ' Porrocaecum anirale (Rud., 1795) . -, -

1845 Characters, Hartvich, '1959: Author Mosgovoy, 1953.

39.0 Body Length 24.0 -48.8 58.0 "" 81.0 - 1.1 " Breadth 0.3 -' 1.5, 0.87 - 1.32 Head Diameter 0-398', -' Ventriculus Diameter 0.714 Diameter Tail - - - -: ý. O.Q27 Cuticle - - Striations 0. CQ2 Lips Inter-labia 1.6 ý- Oesophagus 2.013 - 2.112 2.98. - 3.47 : 0.346 Ventriculue 0.236 - 0.36 0.45 - 0-47: - 0.314 x caeoum Intestinal 0.544 - 0.821 1.07 - 1.48, 1.064'x' 0.153 Cloaca 0.212 e. 232 .". . pairs Proanal papillae ý'12 Postanal papillae 6 pairs 0.643 '0496 0.64 Spicules 0.47 - 0.532 - 0.58 -,--, Female =ý length' 39.0 74.0 60.0 126.0-- 59.0 Body, ° - _ .. 1.7 0.95 2.0) 1.4  Breadth 0.75,, - _ Head Diameter, 0.21 0.23 - Vulva -30. o 8: 7 Vagina 0.43 ` Anus .. - 0-414-,, Eggs, 0.08 0.11 0.081 0.086 x 0.066 .., : - =; '0.095 , x- 0.05'x . 0.075 _sx 0.062 - 0.065 Order: SPIRURIDA_ ' Spiruroidea "Super-family: Syn.: Spiruridea, Diesing, 1861

Family: Physalopteridae Leiper, 1908 ý Genus: Streptocara Railliet, Henry et Sissoff, 1912...

Syn.: - Yseria Gedoelst,, 1919

15). Streptocara'tridentata (von Linstow, ' 1877) Railliet,

Henry, et Sissoff, 1912 Syns.: Filaria tridentata Linstow, 1877

SDiiroptera tridentataý, (Lins tow, 1877) ; Newman, 1900

A- female `specimen was found in the ' oesophagus of the Great

Black-backed (Larus marinus L. ) Skomer. Island Gull, on - July, -1962. The length is' 17.2 , min. and, the width 0.23 mm. The pair . of tricuspid papillae from which the - apecific name, is derived ý ýare,, z ,. (Figure 10.126 mm. from the; small lateral, lips, 183). These -,project,

for only 0.004 mm. beyond the. lhead. and have a : lateral -papilla. on ,. -ý

each side. The' anterior part of the, oesophagus is 0.12 'mm. and . , ,', the. posterior part is 1.049. min. long, giving an, overall length , from tip of 1.169 mn. The anus is 0.188 'mm. : the of, the conical', _ 'a slit in the 'middle, the body. tail -and -the vulva', is transverse of : The vagina is ', 0.25 mm. ',long. ' The eggs', (Figure 184) are embryon- - , , .; when l aid ' and thick-walled, 0.037 ma., by 0.02 mated.: ý. Discussion

is-the'first"record Streptoeara`tridentata'from''the ". -This of Great Black-backed Gull in Britain. Previous records are -the. -, Diver_ (Colymbus stellatus Pont. ) - Baylis, ' 1939, Red-throated _': and the-Herring Gull (Larus argentatus '_Pont. ) - Pemberton, ' 1963. , There has been no redescription'from British material.

Family: Acuariidae Seurat, 1913'(a) .

Genus., Seuratia Skrj abin,. 1916

Syns: Rictularia Fröhlich, 1802 partim; Acuaria Bremser, '..,

1811 partim; Spiroptera Rudolphi,. 1819 partim; Gnathostomata ',

Owen, 1836 partim; Pronostemma Gendre, 1921 `` -

The genus has shipleyi-(Stossich, 1900) as genotype. There

are' four species..:

16). Seuratia puffini Yamaguti, " 1941

Yamaguti' described -_the 'species from the, cardiac region ýof the'.

stomach' of the Sooty Shearwater. (Puffinus griseus (Gmelin)) in'

Japan. - The species has been found in all specimens of _the. Manx

Shearwater (Puffinus 'puffinus, (ßrllnnich)) examined 'at Skomer ' ' Island in the summers of , 1960,1961 and 1962. The anteriorý end . . ,' invariably cardiac lhalf is inserted into the gastric pits in the - ,.', with the remainder of the body. hanging free. The "are worms , -:; -. - remarkably viable and` living, 'recovered from. an r. specimens were . :. oiled bird which had been dead for three' days. Description

Male: The body is'white, °11-- 12.5 -mm. long and=the' tail is ý. ýinvariably rolled up. The maximum width is 0.286 -ý The lateral _ mm. . BLANK IN ORIGINAL plate 49

$euratia puffins yamabuti; 1941 ,

Figure 185 Anterior end. H. P. x 10. Figure 186 Male tail. Ventral view. H. P. x6 Figure 187 Male tail. Lateral vier. H. P. x 6'

b 54

185

O

3

186

ýi'ýýý a lips are not, well developed and. at the ' apex- of the: lobe, is a-

cuticular-projection which has the appearance of'a papilla, rather, -, ' than a, tooth as described by,. Yamaguti. There - are four cephalic

epaulets (Figure ' 185) which 'are 0.074 mm. in diameter and bear-' spines These twenty-eight to thirty-two backwardly projecting ' .-, ,, 0.009 mm. vary between mm. at the free posterior, edge and _0.0185 , 'ventral at the anterior indenture. On, both the dorsal end aides'',, from the '. are found large tricuspid teeth. - Each is `0.115 mm. =

anterior, extremity_and is-of nearly equal, proportions,, 0.055. - the 0.058 mm. long and 0.056 min. wide. Roughly, 0.19 , mm. behind These anterior, 'end arise a-double row of lateral spines. ' are%' 0.017 long the body. length. mm. , and persist for two-thirds of - ', _,, by is distinctive The buccal tube is 0.154 mm. 0.02 mm., and' ,with 'part the ,, a thick cuticle ' and' many striations. The anterior of ° . yt

oesophagus is 0.685 mm. by_0.049 mm" and'. the , posterior ' the 4.9mm. by 0.09 mm. The -cloaca is 0.23 'mm. from the tip ,of blunt tail (Figure, 186).,, There, are seven; pairs of lateral-papill--

ae supporting the cau. a1 alae, four-pre-anal and three poet-anal.:, ' In addition there are mammiform. pair' in', the mid-region just. _ , _a The are unequal (Figure . 187) ` posterior to the ` anus. , spicules --°': and. has flange. The,, is stubby, each ýa narrow ' right spicule, , the left 1.09 0.215 min. and elongate, mm... -' . Female: Except in the uterine region the body -is, translucent `., The white and up to 23 mm. long and 0.34 mm. wide. cephalic 0.085 marginal teeth; epaulets are larger, mm.`i diameter. and the 0.148 from the are 0.021 mm. ' long. The tricuspid teeth, are mm. , _:'

Family:, Ancyracanthidae Railliet, 1916

S. F.: Schi, togkovinae Travassos, 1918

Genus: Viguiera Seurat, 1913 (b)

17). Viguiera euryoptera (Rud. ,: 1819) Seurat, 1913W. '-, 1 Syns.: 'Spiroptera euryoptera Rudolphi, '1819' Acuaria laniuin Molin, 1866

Spiroptera euryoptera 'was found, by Rudolphi (1919) under the horny layer the the Great Grey Shrike (Lanius. ", of gizzard of , excubitor"L. ). The first satisfactory- redescription'waa by' -, Seurat (1913k)_ wvho used the species as : genotype his new genus,, _for - - Viguiera. Other hosts in Southern_'Europeý and'North Africa are, l, '. , .. the Red-backed Shrike: (Lanius collurio L. ), the Lesser Grey'ý, (Lanius Shrike 'minor Gnelin) and ? (Lanius rufo redescriptions Recent of new hosts are-Singh (1949) in the'Bay

backed Shrike (Lanius vittatus Valenciennes)-and, the White-bellied

Drongö (Dicrurus coerulescens (L. ) Vielliot)-in-Hyderabad;. Chabaud

(1957)in the Woodchat Shrike (Lanius senator senatorý(L.

Tunis; Wahid (1961) Tree-Pie (Dendrocitta -in. and in'theIndiaan (Latham), vagabunda ) from Pakistan. :, The, -author has three. - specimens, -a male and two' females from the gizzard of Blackbird , --a. . (Purdue L. ). The in (arch, 1963"..: merula worms were recovered . Description _,.. a'- The worms were slender and: each end tapers, to narrow

ý." extremity. The head ' is' most difficult to interpret, the most description being "that Chabaud '(1957). Because "' adequate of - of ... -',. - the limited material end-on views were not attempted' but lateral :: BLANK IN ORIGINAL lgate50

yi&uiera euryoptera (Rudolphi, 1819)

Figure 188 Head. oil immersion (a) lateral view (b) dorso-lateral Figure 189 Male. Anterior end. H. P. X 10 gibure 190 Male. rail end. H. P. xb Figure 191 Female. Vulva region. H. P. X 10 Figure 192 E99. oil immersion. 513

188 b a 192

1ý

90 89

o_ 3 3 514 immersion There views under oil were possible. are no' lips but --.- a pair of dorsal and ventral cephalic plates ; each' have: tiny- spines

on the inner borders and. a, scalloped' outer border, (Figure, -188(a) & (b) ). Laterally to these pair - plates are ,a of comama- shaped ainphids, and a littler more posteriorly, ', more median pair _a of cephalic papillae. The-buccal capsule is elongated' cylindrical'

and can be 'seen to be an, invagination of the , cuticle 'comparable`.

with, the stomodeum of'coelomates. It is, followed by'a bi-partite-- is the oesophagus,, around the anterior muscular, part . of which nerve-ring (Figure 189). The glandular posterior portion i8'',

surprisingly long, between one third. and one_ half the ' body `length. Male:, 4.5 mm. long breadth of 0.178: mn., ' The with anýaverage : .:: -. diameter the buccal is' 0.025 and across, the;, '', across capsule , ýmm. tail 0.185 The . thick, distinct mm. cuticle is 0.017 'mm ,with - apart -, striations 0.009. min. apart in-the head region and 0.016 :ý. the long, the towards tail'. -,- Thebuccal -capsule is 0.028 am, - and nerve ring, 0.034 =0 in length, is 0.198 mm. from. the head " , .., (Figure 189). The muscular the is 0.347 mm. part of oesophagus. , long the is 2.108 -making total, and glandular part ' .mm. , a -length, of 2.455 min. The tail, ' is spirally, coiled to, the left and because,, . -, there was -only. a., single, male, no attempt was -made at'. uncoiling the, tail in, what is' forBritain, 'an, unique, species. 'The

spicules are 0.007 mm. thick and unequal, -the'right measuring 0.132 the left" 1.165 that is' in the ratio 9` '.. mm. and of .. : '1 (Figure 190).. It, was difficult to determine the post-anal and` , " papillae, those were traced were: pedunculate'ý ý.: .. pre-anal --but which 0.015 mm. long. The left alae 18 larger and wider, and- both, fine the base. showed at .=-a Female :ý The species are 7.8 mm.' and 9.1 mm.'_ ' long with, a breadth

of, 0.221 0.279 Thee diameter across the head is :'', mm. and mm. _ is 0.029 mm. and across the anus 0.068'mm. The buccal-capsule

0.033 mm. long and the nerve ring 0.18L4 ` mm. ' from the anterior end.

The muscular portion of the oesophagus is. 0.312 mm. long but, the''.,

glandular portion could not-be measured since the, ovary, ý,which,

is parallel, overlaps 'much of its length. The cuticle, is- 0.018 -mm.

thick with striations 0.021 mm. apart.

The 'is 0.107 '- from -the tail. The ' is flask-'-, '' anus mm. vulva _a 'expansion 0.03 in front, '(Figure 191)41` shaped cuticular some mmo . The aperture to the, vulva is; circular, 0.016 mm. diameter and the-flask is '0.058 mm. by 0.039 The ovijector is `. mm. muscular ,;. _ ý- 0.031 min. wide and 0.165 mm. long. The eggs are-,: _thick-shelled - d (Figure 192) embryonate when discharged,, 0.036 -mm. by 0.021- mm. ' , . Discussion

Chabaud (1957) and Wahid (1961) did not give measurements are, in their redescriptions. Those Cram (1927), which,, based of , - on Seurat (19130 and Singh (1949), from, new material, are given . _ in 38. The feature 'for=ncomment for =comparison Table . only calling ,- show-striations.; is that in`the male both left-and. rightýalae description left In the of Seurat ' and Cram, the , al a; - only', shows striations, while Singh makes no, mention of striations.

This ' is the first "record of 'the, worm in Northern Europe the Blackbird (Turdus L. ) isy a new, host.: All and. merula . )- _'-previous'ý 516 BIE 8s Vguierr euryoatera (Rudolphi, 1819)'

(1927) Singh (1949) Characters Cram = Author_ Bak Body length 5.2 6.75 3.5 4.5 '4.5 - - ,.- . " Breadth 0.125 0.204 0.178 0.025 Head Diameter - - - Tail Diameter 0.185 Cuticle, Thick 0.012 Striations (Head) 0.017 0.009 (Body) 0.016 a;. "" - Bticcil capsule 0.032 0. C28 0.028., _ OesophagusAnterior 0.370 0.29 0.347 Posterior 2.33 1.23 2.106 Cloaca 0.41 Preanal papillae 9 9 Postanal 2 2 2 papillae _- Spicules Right 0.192 0.194 0.332 left 0.93 1.136 1.165 Nerve Ring 0.263 '0.198. (distance fron ant. )

LeMIL e Body length 9.0 10.0 6.0.:. - 11.0 7.8 - 9.1 Breadth 0.290 0.25 0.273 0.221,- 0#229,, - . Head Diameter 00M 9 - - ,.. Tail Diameter 0.068 ;1 Cuticle 0.018. Striations 0.022 0. C21 ' -: Buccal 0.033 capsule , ,. Oesophagus Anterior 0.78 0. /12 u.> 0.312 Posterior body length -3--l. Vulva 0. Vagina 0.165 Tail 0.035 Amos (from post. ) 0.102 0.107 Eggs 0.032x0.018 O. C98 X" O. C21 0.036 Nerve Ring _ 0.263 ; ', 0.184 _ _

'Intradurtinn - "-. The, survey yielded-, thirty-seven new host records for, °_`, Western Europe (Table B I), , three. species are' recorded for

the first time in Europe' and five-species for, 'the first time

in Britain (Tables-B, b) In addition 2, a& . there are forty- four host: new records for Britain (Table B 3) and 'thirty-two,. host for new . records Wales. Several of the new records for

Britain and Wales are 'due to the, hosts not having been ý examined previously and are not of -exceptional interest since. the presence of the 'helminth `is to be in species , . expected. birds throughout Western Europe. The records, for -Wales have therefore, not,; been-listed. - The writer must that ' it, -stress, _ has only possible to confirm records for- Western Europe . -been In recent, 'years-there have been a great many intensive>-ecolo-'. gical surveys in Poland, Hungary, Czechoslovakia 'and', the',: ý_ ..: U. S. S. R., which, apart" from the briefest' mention in Helmintho-' ,E logical Abstracts, 'not readily are otherwise accessible and°_ a consequence, ; not, 'a few, of ; the _claimod new, host roe ords - -as , . ý- may have been already established in Eastern Europe.. ".ý' . .. S TION A. -Now Reenrda,

Near host re _nrda fnr .teetArn Hur nna (a) Tema.

The recovery of jay erognnum l ongicaudn, 7oonýyýhiaý cýtf Tatum,. Brnehyl_namüs -and ftignntu Q in -theirrespective passerine host requires little comment" since a11--are-established in host host related -species, and, - where known; the intermediate, is included in the diet of all the birds. Only- pý ýoý.^ ýýý. ý 52V Ho Re( for 1.1cst ern : %urone 37 R.,c: at oT ti3

Trcm, Coda = Host. _.., L L. cýroso tura 1oný*icauda (R'ud.. Turdus pi1aris _ Luttrerýa 'urdus L. tionenteron (Price I: clntosh, ý. nusicus, 1955) n tý Parua najor L. - MY ýý rr Paccer donecticus

7, Corvus L.. oonorchis vetio1t turn _ailliet, 1; 00) cora. c

Corvus corona coronc L,

L. nopha]lus delicioauc (Clsson, 1893) Larus ridibundun -. Par_ Nuiaenius (L, ) rorchis rittaciuri. (Braun, 19,01) arquata I'=ra-ronocct)halur Arenrria interpreo'. (L. ) sz=uictricurz Lelopols2: aya, 1552

'ac Lae. (Pud. '1819) Corvus nonedula L. ,,: u- fuscatuc , Lcucochloridium 1927 Certhiae faniliaris L., c, -rthiae I=cIntoch , Muscicapa nt. iýta Pallas;

Lcuooch? 11132, Sturnuc 'L. ori: '.iü. :ýclorýi^. 'zc IiclntQCh vularris

Cc:, tt 60 "ý- a t., ('ud" 181 ) Pratcrculaý ca (L. ) ýtrabothrius cy7 irdr: ýceu^ , arti ;, L.: I'ý? 11 ietina (Skri. ) ýorini (,: cCnin, 1329) Columba oenas, _ 07'1. ' 1370 Larua arc*cntitus `-pont;. jr o , le " otcuc i'riis, T', (I-r", 1819) nocrryptcs riniitus ( 1lnnich) ýarieterotaenia ste11ifera I1ý 1810) Parias rater, britannicus r; cno1e jc (I:. ) ctvlosa -(-. td. ,_ harpe: and-Uresser

Parus i, alustrir ' L.

'Ae ithalus cFiudatuc Le,

Corvus ý'...ýrýýral_si.; du jarýlin1 1869) corlxI. Turduc pi-L.

Prunella oidularia, `(L.

522. in be the, 'are'.:: rvu. _q C or az need emphasised; asý -hostI s habits , rather' different, feeding in isolated areas' and in pairs, or, small groups. Molluscs and insects form, a smaller part of, - their, diet. The ' two ;birds were, shot feeding on pasture at ,, hard-winter the these the, end , of a and proportion of consti- tuents in the diet have increased` '-- Lut!? trema:. may considerably. ,, nnnPntnrnn has been recorded previously only by Mettrick (1958a)

in Hertfordshire and if this species is 'not yet established . in

Europe then the infection of a winter visitor and. passage

migrant-such as Turd us nnsInjin could well cause its, introduction

to the Scandinavian and Siberian breeding grounds arid to "the

wintering grounds which extend to the Mediterranean...:

Pn nnQeaph_ rn yrnr, tricum has been recorded only"-in

Lastern Siberia-where Ar nrrin interpr breeds.. The, smaller. ,,, the and its in the -small intestine: size- of specimens , presence instead'of thatthe the-gall--bladder and , bile- ducts- suggests migration and '.change of internal- conditions, may cause-some, food 'wintering guar- ý-I` discomfort to ;the tr©matode. - The ` in the I-ters; is mainly' molluscs, insects and crustacea, but., in-the ;. _ _. breeding grounds is likely to be supplemented by seeds. -grassy ; --, and terrestrial- invertebrates. If arty= of the former prove

be'_ suitable* intermediate' hosts then-. the species Imay establish itself, in" the resident, breeding - -non- population., Parýýh ýý. lim has been described from of the _nýttý.. members family--Charadr iidao in Russia, and India but only from the, . ',in Rurope. ý'ILimp im; Wir, is family -Laridae ý stn - '1y ' winter _ visitor to Pembrokeshire although there are--frequent' breeding 523 = pairs in marshes and hill, pastures. Thee infected bird was, feeding field in November shot alongside , gulls -in a. marshy where the diet consisted of insects-and their larvae, ' ,pulmonate _- oligochaete, The. infection molluscs and worms.. - almost certainly outside the breeding. seasonwhen it feeds on mudflat ': occurred . s' and estuaries-where the, diet is marine molluscs,. --crustacean and-

polychaete worms. The rarity in other Charadriidae' feeding In

similar conditions, finds its counterpart in the ' rarity of the

species in its more usual hosts, where only two, of sixty-seven

members of, the Laridae dissected were infected with" three and'

one species respectively. LPungth1 ld fern l has been in Mus _ApA. nr nett Ae recorded _t atr Inta in Russia where the host breeds (also in North Norway),.

This bird is a summer, visitor- and- passage migrant., Its food

is''almost-'exclusively, insects and-the bird was-shot in 'deciduous the host Cer ]- conifers where new hi fn nrI ihr , anrifc also":. feeds. The. presence of a--suitable' intermediate host? among the local fauna that the become - ' suggests: species could established , throughout the -migrating range.; 'of its 'original,,. European host over which extends most, of the Western Paleartic including

North` West Africa. vulgaris 'is the first European record, in a- passerine ',bird. The have species is comparatively rare however and-, might ; been, introduced. from, North America where it is common In both Charadriiformes Passeriformos. ; There. are frequent-, and . -r ecords North American land in, Britain,, `;. of migrants , particularly. Passerine the recent theories iriclude, Charadriidae and -and more - 524 jet streams - strong. winds at very high, altitudes"-, which, blow from to the Atlantic to east west across and are -known take large. flocks, off course. One, of the most frequent : of is 1KnIo ta host thias such visitors, "i1 va ri ,a recorded ' of -y; er Charadr iidae include Er j3 10rn spp. while 01 spp.,, ,, and Try spp. The two, new species of Leºin acht orb, therefore,

''Owe' their introduction into Western Europe from either may _ý. Russia or North America. ' _. (b) Ce is - oda TArahnthriiq indrn in FratermAn' 'artie. is cyl -e ua - of. no ' special interest as the species is found in Laridao throughout". ufrtnu and has been recorded from UP In as e and'- f birds -which tend to disperse from breeding pof nn ý . also away tv grounds-in the non-breeding season and feed both inshore. "and "- which out at sea. - All have access to the intermediate host been may be, a small fish. ph y ötvle rot au has- recorded in Lrarns arg ntätua by Linton' (1927) "in North America but. most records are ' in Charadr iidae. ' Gulls äfeed more rarely. on the - , polychaete, worms'( the intermediate, host- is N rei4 d sicn1or) and the only earlier record in Laridae in Britain is by Pember-1 ton (1963)' who,, found the species- in five out of one hundred and zdihundua, The forty-six Li 1is -. single immature strobila, indicates that, other, factors, besides rare contact with the intermediate host may revent, the species from beeomin 'Laridae. established to any : significant -degree in the. gno1ap _s is found in the Corvidae , but ., y1n commonly ý, has not yet been recorded in the Par idae. '-', The ` food of this =ý-ý 525

latter family consists principally of ' insects, spiders and ,. _ of the, centipedes and only rarely earthworms, -or slugspý more'

likely intermediate hosts. The occurrence' of Aplop., jrnkgjq, , duiandini in modularise Trogt od ody '. inel a _trogl and Ent n lb n-ynrr is-. unusual as the intermediate host, is =

an'earthworm and the, Wren and-Pied Wagtail feed'chiefly on, insects.

(c) ? mat cA_.. ü inflate The intermediate host of ý; cri ý.^11 can be - common worms one of several oligochaete and yet the nematode.,; _ not has previously been recorded in passerine host "'although' ,, ' ,a diet the worms- form part of, the of many of the perching birds. " The in Tur tos- meru? is single occurrence a, quite , exceptional and is ind is ion that' factor host- if is ity : an' at some of spec ,-must have', e been negated on this ' occasion. -'Cf? 1 aria or is very=

common in passerines and of the' now host records only, Anthij'n'l

's noTh need be mentioned as,, animal matter, constitutes 'a',-'

very small' proportion of the diet. The record of Capili in C i-q f1 is is' of interest for the spp. , ýrduue1 nvirögtr same, ° D., Wakelin has that the be reason. 'Dr. suggested species ,may ,' C. -nnl (from the Chaffinch) or 0*1gid (from the Hedge;:; Sparrow) but. inadequate original .descriptions. make idontifiea=

tion- extremoly-difficult. Ur-. yi tiiÄrý ew has, been recorded` from the Lariiidae:

in Southern Europo but not from any, other. - passer ine=family. _ý ieus tarry is The intermediate host-is Liimhr .ria which ° also Part of the. diet of many_,:other''- Passer Ina s. '-,, The Laniidao 'breed, ; 52G from the Baltic down to the Mediterranean' and= it is surprising

that there are no other records. This unique infestation, is', evidently the result of ýa, break-down in host-specificity,

2). T' iae ra. dd for the firsts in Enro pc cm .i ma ;

(a) 3.m tnt A.

The occurrence, of 2r arrnnaceph nng; CrinPtri(! tiri and, Lp=-

nhl oridium th ba hai- been discussed above. ( b) NAmatad,

Snurntia, puffite was described in Puffmus or Inn u from the North Pacific. This remarkable bird nests in New Zealand,

Southern South America migratory is and and fits range -,well - beyond the- in both the 'Pacific Atlantic , equator- and oceans. It the coast North appears first ýalong_ east of AAmer'ica. 'and, -then',, Greenland and only reaches the European waters as a', scarce

autumn visitor for, its, migration.. Its principal -pre-nuptial _ ,:. food is the squid but smaller fish such as sardines often - are , i taken. ' The species was recovered in Puff I us pff, nn.R which The breeds in Pembrokeshire. parent birds fly. daily south-_to, `-',--

the Bay, of Biscay, for sardines with which to 'feed the- young.. ' and further in September, - when the schools move south the, - birds follow and may even winter off South,. America.:: - (One bird.:

ringed at Skokholm in 1952, was recaptured , below Buenos Aires 6,100 time miles). At - this of 'year, 'it -overlaps, the northward ,` PutInns us cross-infection to migration of . and appears f . take place. The-intermediate host is presumably the sardine. ý. -Several birds, 'examined but not infected. young were _were

Timon-David (1950) recorded Cy clc en1 (Hyp ýaem,ý; ý) in din, mutes Pic pica France. _: The morphological signifi- . the different location, in the host be discussed., canoe of . will The difficulty in explaining the : is due to -below. , ýýinfection the host's being-a non-migratory: species. (Dorst, T 1962,. p, 261 ' If there describes the Magpie as sedentary') . is no _migrat ion', of the host, then a possible explanation for -this unique'- infec- intermediate, host is tion is ýthat the - migratory, -which would, being-an insect. Alternatively, the of course limit it=toits . infected droppings, of a-, migratory , bird may have caused 'the, ý

infection.

The record of U11Cn oridi, lm, meI os i 7f is discüssed, -hi _.: above.,, stý, rn i vu nr 4 is a -winter -migrant to Pemrokoshireý in Northern and-the infection, may, have occurred Europe where = the species has been recovered in Charadriida©. (b) aastgda ..,... na to has, been recorded, from insectivores _g nog eis. -LH _) ,- in'hurope- but not previously: from certhia 'rnm13ja hrIt r nnniR

in Britain. This-host is non-migratory and again must S-ubscr'ibe, the host-i, to a, migratory- insect as, -intermediate the infection. Agma was the : original host for,

_. Ftm ia" It undergoes', 'winter migrations ands , rtarfadea _irý . . those breed ' in Britain reach. the west. coast., in exception-: -which There was no means of deciding whether ally cold, snowy, winters. ,- - ' 'whether', '. the bird examined in February, 1963' bred in Britain' or. 5.2 it was a 'migrant from, the Continent. (c) Tema t oda .. of' The occurrence V gn bra a iryop in iird is r,ýer, u1ý since:. 'as in Northern Europe is unique is mentioned- above-it- in the family Laniidae. Only Ln A a normally occurs l «Q s, . mr ýý is an annual visitorýin very, small, numbers to the West Coast .ý . of Britain, ' particularly the South, Irish Sea and the. Pembroke-' This bird, breeds in Belgium Poland shire coast., , and, 'and` winters in tropical West Africa. (The_ species has been recorded

in this host in Tunisia). -It is considered that rather than°

this record: being a chance 'infection, of- a resident, bird, the There Blackbird. was a. visitor. is a-considerable mig- -itself', Blackbirds, breed-: fr -ration of Continental which 0M_-,,Scandinavia, to North Portugal and east to central Russia, into 'Britain in the beginning late September, which remain-until of, April. -'The ''- worm 'wasrecovered in the exceptionally cold 'and wet, March', °

1963 when the influx-, of migrants was especially , marked in'- Pembrokeshire. The bird was 'almost certainly infected atits

Continental breeding ground. , Naw Ho-qt rPrnrdy far Britain.

(a) TrP ' ^ttf1a and . P_at04A ....

There is little, interest attached. to the-new British-

records' established hosts,, in Sur ope.:.: In certain asall-are : ,.. igin'al' the', new hosts =listed in Table'. B 3, are', the or --cases, , ., were described: hosts from which, the species - hi

rh igndaZa Diet z 1909)in N.S'an na ^rnaa-t^om'(L. ); Cy-7ocº'ee1ain (L"); rin CC. ) mutable in, Qa___1inula chhpus. -Tatrabnt J pia l for Britain Fort: '-fowr n(! Nowt (ecord0 -, 5301

Zr e t,ato01a iIoct

"'lc--iorc? (Ruco1pI'. i, 1302) Hirundo rustics 7. " xio =culocuc _-- 1c, jcau(la (`ýuclc1ph 1309) ýor11uJ corono cororo L. _erc onuni nc°. , (L. '), ri rt pica Pica

Iýitýtrez: (Fricc L, iicintosh 1935) -ritlýacua ruiýeeula no1o h1oaý.: ia nonenteron xaartert 7, 1900) Pica ica (L, ) oorchio petiolatun, (2niii. ict s ; 1393) Laruo L. ` yr: ;o ha äuo del cioaua (cls: on, t arinus '

Inruo co.nun T,.

Corvu. -carono L. cLino torn rcyoluturi CErtilicl, 130') corona `_- Coryue3 rý ýr nonedula

ttuncniuc CL ) ýcxýýýtl. i itc"ddxnýý CDict: '1ýCs') arctuatc. -" a rý, , e ., ,r« Numeniuo phaeopus (L. )

ß'^ 1819) Coryu. fru 1uL . ac ýIýez: ý:o ýuýcbtt. a Cýý:týolp , o Corvu ; corore corornc I ._ (Zeder, Calrula. loropun "(L, ) Cclococ(wc±ococlü:. ý) rrutnbßi. c cf;. 1800) r=u1icttra u rr L.

Cc"`, 5_ -0toda - _' ýetrAbot? iiýuýr. ý9) Laruc m rinu3 L. iriuc cylý . ýu'raceua ,1 rr ýý Laruo tuacua L.

Totrabothrius (LBrinbCrt', 13 0 Laruc riarinun I,, '. ý. erootric . Poclicepc (L. ). Tetra bctl: (. tbild card, 17'0) "-. riu . ll. 2riorinu:. l' _crictatu's b3leri 733) (%hranyc, 1 vom Corvuo coraL. unOula , _ 1'arictcrotaenirý 1669) Gä11 na o ~a11i-. ýý . erýüro Krabbo, "" ý'ýo ( Fuhrmann, "urduo mcrula L. " : notaenia urir_ oronata A`1LtS AýA% VtlS ', '? n (! {rat be 1136 9) e1? nC, CP. Oý E' }S (iý 1 tiý:? aif ^ , ,_.. Ifulica 'atra L. 531:

hrieno? 1927) Inas L. exýis (ýI. ) ducts Iio (Linton, nlutyrhynchoo

Sonateria molljociria (L. )

II 1 a7) : 'aruo. ma jor L. uenolý nis (H. ) ýýcir rya ( uizr"auil, ß

0) ! arur L. (. a1 I9 iur enol Pi (N. } ,, <

Aeritha1o i caudatus' L.

Il (ächrünl: 17 0) Corvuc corax L. mole»ia { }I. ) ^erneýtitu2ua 1 '". Liorchis Fulica L,....,. acualinäta (C1erc, 1902 atra Anlonaralcai;, (I: 1 9) C 1idrio Innut 3 ý:rachynhal2om rabbe, T inlas ta"Luaus"T., = 'ý`ýýýlo?ýaralsirý ýý 1är1 ((3roczc, 173')

Aý 1oTi4 (I: 1.69 Qallinaro r7allinar-o' L. '- '. rak$iý hirýuta rabbe, . So: ia: 'i (L. ) (Puhr. 1ann, 1913) ýateria i.to1 a

I10 atoda

CQ- ý'ithaeu$ rubecu1n rielophi1os i11aria cortorta (Creplin,. 1339) 'art ert; : nas rlatz-rhy rchac L. ;.

ITuIlcniuu arqua`o (L. )

A1ecuda L. urycnrý. c -

C4ritr-ýCnecurz' (:: 1309) ',Ica torch L. : ýsicul; °erurt udolpllli UrEa aalge (Pont. )

Puffin tc nuffinü3 (Drunnich)

Lamp" fuccus -L.,

town 187x) Lcrucý marinuo Z.,° ` try Banta' on ?tin: . _ntciearä a _ u.. 532 QY.1InrirrannP (Rud., 1819) in Larti 'iannUi L; '--Tetrabn hriii . 0s (Lönnbergý 1889) in Latu4 r inq L. pa a -at., nniA mhr, y Krabbe, 1869) 1n Qa 111 go ra11. (L Chn,, notrani In 1T13rd nninarnna (Fuhrmann, 1908) i meril1 L Hv, nal an tn- .a .pi (H. ) ina in. E-^=rtýa orrL.; and Difýrehia a (C1ere, _trt "llrlitta "ý

1902). 'in Fu1inn atr= L.

(b) Nematoag

The ': finding, of, G nlllaria'spp. in Al uda ar rp Qýý'is the in the Skylark first record of any eapillarid worm in Britain. -, stra is 'a rarity in British birds, rk-nir8 .r fdpn .ata -also and in X01yrnhil the only previous records occurred 4 ata1 i at and , Lard

ý b TIONT T TnnidA r) helminth sn_ A. _ "ipa -n Over five hundred and sev©nty-six' birds belonging to examin©d. The twelve orders and thirty families were total species twelve number of bird- was seventy-nine', of which , were. " Three hundred fifty-roar free of, helminth' infections. and (61. f%) with at least. one birds were parasitised species . of group the hel'minth. The most common helminth was cestodes, . -: forty-three species and-two-hundred-and ninety-four birds were most group Parasitised (82.2%). The' second common wer4, 't he' trematodes, species `and eighty-six birds were para. -eighteen infected sitised (21"x+%) Finally, eighty-five birds were with ". _ . (20.2%). nematodes seventeen species " d to host 1) of holmint elAsaP4_relst a rn end fmi1i . birdsinfected in the survey, is shown, ý_ The percentage of ý -, ,ý , _, 533 { 4. in-Table B When such disparate-numbers of `birds `for each

family -were examined, it is invidious to draw too many conclu-

sions concerning the susceptibility of family'. groups', and orders

to helminth infections. Certain general statements, may be made, while admitting the tenuousness of arguments based on 100%

infection rates when only, one bird dissected. was .°, The incidence of trematod©sgenerally is remarkably ' low.

The papers of Lewis (1926a) show a far higher. percentage ' infe s- tation in marine, shore-birds and waders. Davies (1937 unpubl. ) ,' subsequently recorded fewer at Aberystwyth and Williams (1961), _` fewer Recently D'. Gwen Rees has that the again*' . stated incidence of ' parnrehi i pittrciti'm in the Lariidao, is a fraction: of its former, prevalence when she did her classic work-, on the ' life-cycle. It has not been possible to ascertain the factors , bringing this scarcity, about. The occurrence, Iof-Himaathin app., number is as rare 'and the recorded in the Sc olopac idae ,=is :,..: , supplemented by Gyýrhýýlus nu _nii sp. nov.,. in a migratory bird, Ni Argwit . Resident, birds, like Hr mntnpus, ogtrn- , (Family 18gus Hae mot op idae) are st ar t ingly lacking in the, -

species recorded- in'other parts of Britain and Europe. forty-four Lariidae infected-by in - were all Gymrrnphn11 s dal it i ogu., ommiting this species, only three, birds were infectedýý-' two. Paroreh_i i- tac1 1i, Hiºn hir+ The with ß and' one with :1t Qiongnt . intermediate host of G_' Ani ieiaa Iq11 is, unknown. This is, plentiful on, the c oast and in some way the' spIor ocyst and' r ediae of the species must be more resistant, to detrimental factors than ,are iN 1 N1 1 Ü 1 1 N CY M "- N r-1 r " 1- CV rl ,

h 1 1 1 11 1' N iC : - - 1N 1 ý 1 .. 4p 0 U1 M M NH MN1 co ri 1 1.

ri ýD H I1N H 'UN ri ri 11 1 ýU rl v f N N M r' 111 - " r{ r r-1

ri ri ri Oý 11N. 4Mr1 1111, r 1" CjVNN "-f f` 0.1 e-1 .ý1 rl

I1N1 -r4 1 ., } 11 r4 r1 iN r0 1 11 r-1 u1 º-i 11 r-1 1 M1I ri "n . -1 -ý wi

rl 1W11 r-1 11N1ri 1 N CY q -* 1 8 e-1 1 r-t 1 i"i 1 o 1 1 N 1 cc) 0 v ri ö g fi 8°N'M°0'-\ SS8S8 ý= M ý' ö g° °ý.

cv ý-1 CV r1 O . -i NMN+NN r' cV TN ONN ei n NN a- Iý 1 fý r-i .-v " r-i CO ºi rl r-1 ri V- N LC%

cm NMN r1 MN$O qN \D 97% cli "- -4 u1 'D f co co N ri P-4, Nv " co i rl -t ri N1 rf ýý t" r-1 D +i .N

ºý1 CO ri ý-1 rt tý N ý-i . P-1 9.4 ri " VL "G co ult M fý NN rq H rl rß N ei A d r-t 0 I 41 4A .Aý. ýeý ö Cý Ami r-1 tl 0mU0 0 1r VdäH to raj ýº 'A

C.) cn

wi ýi . Q c7 n O -mom nn0 on io'i .opp

1. ..-4r C- Ell pq ßg1ri -,, r4l C -w AvS0 k a Poo E 535 those of other trematodes. There may beý lack of lime ' in . a river, presence the sea and in'the, principal - the C'leddau. The- ) (shot the of Cy- ocoP]. ý(C. mu "abi 'in the Rallidae on" Cieddau banks),, ' this, -does not necessarily contradict since their diet =also includes insects and larvae, worms, slugs, and tadpoles.

Most birds in` the of --the ocean-going' were examined summer when previous "surveys have shown seasonal incidence to be at

its-height., The majority of-the Charadriiformos_ and'Anatidae= taken in 'winter incidence is were - when seasonal often lowest. But any, seasonal, incidence was not so marked in'. the cestodes

from the same. birds, and . the - attention is directed to the different to, the intermediate 'host ' -invertebrate -phylla which ., belongs.: Trematodes_ generally, use molluscs and test odes use'

polychaetes,, oligochaetes and assorted arthropods. '; 'In' addition' to the' absence of lime in affecting the molluscs, one -might

consider, also oil- pollution from the tankers, using, the,, landing,,, the the: stages"of new oil refineries at -mouth ý:of, the Cleddau, the sewage -.outfall" from coastal, towns like Tenby, , which make . ' most of the, local mollscs inedible and, finally, and ,perhaps: - the increasing insecticides '-'which' most significantly, uses _of inevitably get--washed-into the_Ceddau and into many--fresh-water ý:

streams. which flow, out from the cliffs andacross beaches. °.` -the The-, sparsity of -trematodes in passerines points to, a lack: ot the, cause.. ý. ýDespite, -lime as. _prime the, variation: in numbers for each family, It As

possible to, draw, some conclusions -.on-the pattern'shown- when, 536 relating the host group to their diet and, indirectly, to., ' intermediate This the availability 'of a suitable _ho'st,; is'-

to be seen in Table B where habitat and diet are related from helminth to the ,,number of species recovered each group. A marked-increase in infestation is shown in all helminth

groups the source of infection changes from' aquatic_. as, , _inverte-' brates to purely 'terrestrial' invertebrates. trematodes in birds' There are many records of aquatic ý and their absence' from this survey has been, discussed (above. " The;,.,

trematode; infestations increase as the-diet changes toýpulmonatG faac, ti1a snails, *, for Br a hYle 1 when slugs, are the or as _M1In , intermediate hosts. The high rate Of infection in the Turd idae the inclusion of slugs in their., appears, to be - due, to and worm diet. " _.. .". birds have Generally- the , which a predominantly- veirtebrate,, than-trematode diet have more nematode infestations or cestode group f) Strigiformas. Similarly, infestations - group - a) and' °; life-cycles than trema-;, nematode's with direct are more common todes in birds in which seed-eating - predominates (Passeriformes;

Prunellidae, ' Fringillidae, Alaudidäe) and "where vegetable matter ' -. (Columbiformes; .is equal toi or in, excess ofq animal matter -' The twonematode Passeriformes-. St urnidaePasseridae) . specie i in ( marked" with an asterisk) ', both= the freshwater group, . occurred" unique in in t=hX04 ' is " the group in that it 'An.43 , ni ..which ° flooded pastures. The inter= regularly feeds on, marshes and. = concerned mediate hosts for the two species', are earthworms, so ..;. the final host comes into contact with nematodes not normally ;,`.

4A N

09

Pr s

hO r4 1 Iz 1- Kl ed, OOý

Ä

qj O ýM ý 'd wý 'w 'ý .h O P4 ww o1 co cli di ýP 1414 Oý a IIýi1 Oh" mýJ "O ,N Id

of CS wVw90 qw w wS O fl yý [ý. 4-1 0., o 4 'M 4J :i. 4 C) 41 qwöi: d. 14 Ow +1 O e-1 aq .. 9 .O N4J ßßh d@ 0dOp 41 - 4,13 II ýº '1 1v - ýv Opp,. to p W .Ovü

. eK

n °) O,. Al n le n 0% w ý C min t

C20 .00 04 4, wU

I% 0 4b ti P% !Aý4 t (D o

E-41 IP, 539 % occurring in freshwater, birds.

The number of cestodes increases as the diet of, the birds changes from aquatic, insects to purely terrestrial insects is its highest. in birds - 'as the _and at arboreal such Paridae. The insect larvae must become infected and, the

cysts survive- metamorphosis, - to infect-- the birds. 110 from io th 2) " sPOIria no o-1 m warrp nv red.,

The,, twelve species are listed in Table ýB 61 together with

their diet. ' A single Tndornä t ndorna was examined and there

are, in fact, many records ofhelminths in this species 'in 'which Britain. The diet is a mixture of marine and terrestrial'.

constituents, is one, from which many infestations are to be The bird was in in January' "1963 the', expected. shot snow and _ absence of, holminths-, may be an indication of 'seasonal''variation., The bird is rare in West Wales. similarly, one DAndr examined. The, only lul-q-^ý .r was >', bird is locally common, in West- Wales- and the 'diet indicates'=

that not many helminth infections are to be expected as,as ,,, principle food, wood-boring insects, are unlikely to 'come into

contact with the droppings of othor birds. The only record, 'ý :_. has traced is (1937 ) the writer Walker unpubl. who found, ':`:, :". intin. fragments. 1939 2. ätüm III spp.., -(Baylis, records, 'petin n ', Chop cratorifamis from' the Green Woodpecker. and which ,-; eats wood-boring -insects and -ants. Ants dare more suitable" intermediate hosts than, theinsects) . The remaining birds-are: all passerines. A single nar he

oenanthe was dissected ' and, the 'diet' almost entirely insects 5 40 T,Aßt Si; Alist of Airok species in, whic}x no he1riinthc tiwcre found.

Crdcr ; '.. peciec Diet : , r

Anr>eril orm. en Tadorna tadorna (L. ) urine riolluoct,, smal1, cruc3tacec; zso! e inaectl . ona11: fish, norms 9 veg

Piciforner Pcndrocorun L. - valor Wood-boring-incecta; 'I- Fartily llicic (Great cpidcrc; seeds, : ae - spotted wood- berried. _nutc, pecker) Paa^eriforriea Oenanthe oenanthe (L. ) Alnoat` entirely.. insects, Family: Turdidae (: Jhcatcr

Sa:: icola (L. ) Chiefly corze. tornuata -insects;. (Stonechat) spiders and earthworms

Farmily : ; ylvidae Pllvllooconuß colt bits Chiefly. incects 'Jielliot (Chiffchaff)

ReC. Fariily: ulidao gc: gulu3 re r°u1us (L. ) Chie fly' in: ectn and , (Goldcrect) spiders.

Family: Fringillidae Chloric c?:loris (L. ) Zeedrsý berries, Buda, beetles; __ "., (Greenfinch) some young . partly on inoects.

H It Carduelis c. britannicus Chiefly aecdy, some Ilartert. insects. ''--' (Goldfinch)

n It (i.. Carduelis cannabina ) Chiefly useec'a (t ;e c4) (Linnet) some incectc. '

ft lt Carducli flarraca (L. ) , Chiefly ýecdo ý co ýa (ý, minute. inoectc'. and ýýod?ioll) larvae.

It it Pyrrhula (L. )- Seeds, p rrhula -berr±ec, in budz. trees) (ßullfinch) uprin j, Young on cater. i1ar . . if H Emoerinia citrinelln Chiefly _corn, berries, ýraos, sorer ; (Yellow l: inrectn,, a ner) _ vpidcrs, 0t but `of . with small quantities ,molluscs, centipedes and spiders, is one from which infections are to be expected. However, the

only record from these birds in' Britain, is given by Williams

(1956 unpubl. )- who mentions Acanthocephalan, Prna Xn us , an trnnsver. 4+;y from the E. A. Lewis collection Aberystwyth. , at ,, ( Madsen, 1945, lists C. c ontortn, which has oligochaeta , an intermediate host, as a non-British record)

feeds mainly, on insects but earthworms are included.:; There are , no, records, for this species from Britain. Ph;jI n-L- 0 hit s andRegý1 pr 1u are also insect-eaters from which there

are no British records.

In contrast, the remaining species dissected, which belong, ',,-

to the family, Fringillidae, are principally seed-eaters and the'

few records are mainly Ci ar. a spp. with direct, life-cycles... Wakelin (1963 ) found Q. i in Ch inc, hI_nr ia, ' unpubl. . - cbn snit or Evans (1938 unpubl. ) also records nothing in Qb11or is or

EmthP I!, ihr n 11 n. Clapham (1957). found nothing, in h Car u innznnabinn. The Goldfinch (Carduel j rar ße1 is t.. and nnn1nn Hartert) , the Redpoll, (Gard 1A 1-.q fI amrn L. ) ý, . _the Bullfinch (P L1ap yr. rhu a (L. ) appear not to have been included in' British ýý previous surveys. - M C. Snn Anni orical eon idrhratinns.. -542 rv

Intr=: nttn. wrote,, Noble and Noble-(1961) "When a parasite is. studied";, ' by itself, only a part, and often a small part, of its 'total , biology can be understood". The total biology, is 'a, complex ,_. resulting from the interaction of the phylo- genet ic tendencies'"

of the parasite, the micro-environment, or the physiological ,', condition and extent of parasitism within the host, and_ the

macro-environment ör host's external onvir onm©nt. '

', -It has not been possible in a general surveý study any ..,, . -to in detail. To do it be ecological aspect so would 'necessary - to examine largo numbers of a single host species (preferably-

non-migratory) over a long period so that-as many features

as possible of the environment are known; the diet; of the species at all' ages, the physiology, flight range, habits. and the reaction to climatic factors would have to be " studied in'

addition to'the parasitic fauna. - Physiological-studies'would .-ý include pH changes within the intestine and the general, body temperature of the host. prepare For each holminth species, it'would be necessary to

a standardised set of morphological characteristic's" -possibly ý., based on computor analyses, but before cbmputor, analysis can', be the techniques of applied it-is essential to standardise methods preservation collection and materials and used in and'-:. '; - Further techniques for histo-chemical differ-'' study. staining ; , , entiation be' Once=-the' morphology of a 3peeies must developed. -, , 543

is standardised, the specific growth rate' can be determined . .ý and its variation according to dietary, changes `and the presence

of other 'parasites. Finally, the range and' distribution that ' the known intermediate host, be -within .of should annotated. -In default of any studies of this type, only the most.

generalised observations can be given here. The only birds, dissoc ted in the Starling (200, suitable numbers were plus) ,f (34) the Great Black-backed (33), the Woodpigeon 1 gull the Rook (21), the Herring-gull (17), the Skylark (12), the

Oystercatcher (11), the Jackdaw (11), the Turnstone (9), ' the ,. _ý.. (9) the Common Snipe (8) ,- Lesser Black-backed gull , the Carrion. Shearwat (8) the Jay (8) Crow (8), the Manx er and . Unfor tu- nately, the Starling, Turnstone, Curlew and Common Snipe : are ` winter visitors and ecological observations such as - seasonal -; variation of parasitic fauna are inapplicable. ' The ' Skylark, , breeding bird, be this-,, although a resident, must placed with - group because ten of the twelve birds were shot in part of-, a'

massive Continental migration during the cold winters;. of 1963 'ý The remaining two birds were and, were all negative. ' each-; -'ý parasitised by a single Cr Hari' spp. In contrast, the;, Mama,

Shearwater' is a breeding summer visitor and for similar reasons

seasonal variation cannot be stated.

The, other birds listed above were examined 'at ally-seasons .

of the year and ones or more fledglings or, fir st-year birds:.. were post=mortemed 'in addition to adults. ' Also all birds , '-.: in were taken in or near their normal, breeding grounds the summer and at points where they normally congregate at other

seasons. -Thus, it Can be assumed that, the diet was the

normal one, appropriate to the season of the. yeär and that. for maximum possible infestations were-obtained the species

in that area. When so few of the intermediate hosts, are known, the they constitute in the be proportion -diet cannot estimated and the total ecology of any species remains an incomplete patchwork.

1) Th the isr°.,. anvlarmn_ . nr_a_itc rn , If the parasite can establish itself and complete its development in host, then the factors" a particular which ..most =` influence its relationship with the host are' the host's' diet,

the age of the host, seasonal changes (including changes "from'ý- to the the year year) I presence of other parasites', and location within the host.

a). Thp. dint: nt` t ha (i). 'extent' The nature of the host's food influences the of parasitism the type ofparasites presentg. was discussed ' and =as ,. Table B 5, above in'connection with whore definitive hosts food shown to be parasitised which - on other vertebrates. are ' by A fundamentally similar-, diet ©nable=-a- - fewer species. will -, ,_ with 'a' range- of specificity 'to parasitis'e' parasite wide -hosts, _ orders In this 'survey, Ilý1äp 14 from: different. of birds. - ý... is, both, Passeriformas and Charadriiformes. ', ý recorded. -from s ...: do not, increaser the- range: Conversely, similarities of diet of ., E' hosts in a host-specific species such as Kymnnn Apii (IL)

f_r ýi ri nga- found only in Qrru unr ,1 ,i

The more specialised the diet, the more restricted the

number of parasitic specics, because 'of the limitation'-to a

particular type of intermediate host. This does not mean ý

that" intestinal chemical and physiological conditions ,, necessarily prohibit the establishment and development of

unusual parasites if their intermediate host should -intermit--;

tently form part of the diet, as is shown by the recovery, of: ,

-HX n1 Pp i (H. ) styl a in Paridae in this survey. The rare infection of these insectivorous birds is obviously 'due to

the small part that non-arthropod matter constitutes in the'-'_

diet, not to any specialised chemical conditions resulting

from the mainly insectivorous diet.

(ii) The eff: hn in the d. the ti . nnt nf. 1 nCn t eri p ra si a, .. Many bird species change their diet according lo the this is season and especially true of the Anatidae' and Charad-, ý

-riidae which migrate from a semi-terrestrial diet during'-the

breeding season, to a purely aquatic diet, whether marine or

fresh-water, at other times of the year. Amongst,, the passerines

many of the omnivorous birds change to a predominantly carbo-ý°

hydrate rich diet of berries during the late 'autumn and early, --

winter months after which the diet becomes loss rich in than carbohydrate at any other period of. the years until the ,`, late Spring. Those in changes- inevitably result'- in a' drop the__-:_ number of species :present and the extent of parasitism.: , 54

The most significant seasonal changes which can--be attributed directly to a change of diet occurred in the three ". -, members of Laridae mentioned above as having been dissected in adequate numbers. Twelve species of helminth were recovered but for the isolated infections Herring,, ý; except of single ,. :' : Gulls with single incomplete strobilae of ophry 1e rnteU14

: arid Disc n! iindiiln in the Decembers of 1960 and 1961 respecti-}' voly, only two species, c1ymnophr 1i ti s dplc n" a and. TA. ä.. höthr Lin ny rnrAg were recorded in all months of `the year.

Even for those there is a decline in ý, species -numbers -present in the winter months between the end of October and the end- of February which is less strong, however, iniAi rinst months The complete absence of the other costodes during-those. is almost due to dietary influences. All three, '_'of certainly ." these birds spend much of the year following the plough when the diet is restricted to terrestrial invertebrates.: At time is this life, °as in the . mode of maiptained so consistently winter months when whole colonies desert the cliff-side°, brooding -places and roost at night in pasture. 'The change, 'of diet is one which not only precludes the marine ; inter mediate, hosts so that the infestation rates are least likely t-obe maintained but, also in decrease in in the, -, results a salinity -, . diet in the,. intestine "the and and "those chemical changes are, most probable cause of the complete, loss of ©stablished in-ý" ýýý'=n , tostinal holminths such as Ana atrnnin"miernrnnt which'-is found in approximately 90%, of the birds examined- in, the ... summer,"months. '547

The only comparable reference: in the literature` isOliger,

(1950) who, recorded dastrobilization in Tetraonidae in Central -

Russia as the diet becomes, increasingly lacking in carbohydrate during the winter months. He suggested that while the birds survive on stored fat, the tapeworms starve through lack of. nourishment. A comparable destrobilization or loss through' the lack of salinity in terrestrial food may be worth investi- gating.

b) The he hca . se . Most authorities have listed the incroase in. parasitic'

species with the age of the host but there are contradictory, reports as to whether the intensity per species increases 'or-. ' diminishes. Where there is a subsequent change of diet, as in adult seed-eaters which include animal matter only, during, the., nestling and fledgeling stages, an early infestation may, die be diminished in intensity. The out altogether or single' - occurrence of Hy noiep above, 'in a first-year

Pug11I noniAh4 is a typical case of- a reduced juvenile;, infestation persisting in an older bird. _ In the gulls examined, there was invariably-a'greater, intensity, of infestation of parasites in first year birds...: ý., than in older ' birds- in ' Himnath_ all cases except-the cestodes., . A 11 " ne, einngnta''and -PArnrýhj-Qp f11e, were ,`found only in juvenile-_ý and -birds : P. -; täc" ium iný particular `limited't o. the `ý _is -younger_ , birds as. the bursafabrisii is subsequently lost.lost.,. Only r, day jn 4 'th©" trematodes limitation' --- oat14 . of showed no age -, and ,,= incroas© in, intensity of infection with age:; ýý an __ ,.: 548 The highest infestation rate ; with $rnir uifr in' ý the Manx Shearwater, was in two young birds. leaving the _ .. , underground nest for the first time. The youngsters, which fed the are on regurgitated stomach contents of the adult - birds, would appear to be infected both by larval- stages, from

the semi-digested sardines and directly by dislodged adults from the the bird. The stomach of parent records -of Rniii, minn (akri. ) boje came almost exclusively. from first, Woodpigeons. The initial diet animal-, year contains plenty of - material and this is replaced by grain during the early autumn':

when the infestation rate began to drop. With the" onset of_, _ the diet in local birds is to -winter, virtually restricted acorns and the infestation died out completely. Either -ýagG or acquired immunity prevented further large infestations of

Woodpigeons after their first year.

All the trematodo species in the Rooks, Jackdaws. and Crows.,,

occurred in the first year birds but the cestodes at, appeared . Conversely, 111 ir all ages. ga: nr seeti was rare' in the - younger birds but in the Rooks at least it increased-in intent-ý

sity as the birds grew older. ,: ý- - flE3nsonal hrnggn. c). . =I ,.. These result not only from the ' change of 'diet and ' habits , as above, but from ' discussed also cyclical changes affecting'.:, the intermediate hosts, hibernation. Most, birds, ,such as - were , examined in; the' winter, months, due. to their status visitors or from 'a desire 'to shoot outside. breeding season. " [the 549-1

It is not possible to 'give' figures-for 'seasonal changes- in. '-' these birds.

A distinct drop =in the infestation of gulls with, , , dei Iel Iagiia was observed during the winter. Whether this 'was.. is correlated with the supposed lack of salinity, which `..

postulated for intestinal holminths, or whether it is due to

some innate rhythm in the relationship of the parasite and

host, it is not possible to state. Dila 4ii ''was more prevalent - in the passerines during the spring and summer a,, ; -,--'fact which cannot be related either to the hibernation of, thee-

intermediate host or to its absence from the diet during the The is true Hyrnnnin; (H. ) winter. same of jln rD_n . >>u_c and again some innate rhythm independent of the host's, age,. or,

diet is postulated. d). Thn i i ty i fa i thn ntp n4 of _t on ini Winne of nth, r jýnr"gai .PQ.;

The intensity of infection and the presence of other= has parasites been reported by many workers as causing. a dec-,

rease in the size of parasites present. This, has -been, observed

during the present survey in Sturnu4 vul C 14 where, the. presence ine'srall;. of, a large c`estode such as Di? pa undid i'has resulted

measurements both- for Il_ t ndO itself mother eestodäs' a and' . present, for example API nprr;, kei a dn jar i i;, The observation ý.,

that it is the-length and, breadth of the, proglottids which'' :.,1=

decreasel', not; the actual number, is confirmed'here. ' The,.

morphological changes affect only the strobila, - the-, hook, '-:s ize and and, shape other features of the sc olex remain unchanged; '. -, . ` does eg& size as , and' shape. 155U a). Change of oeatIon within the ho-qt. - -, ; Only two new locations are, -recorded in this survey,

pronnenphni -i gy m-lcitr incum, in the small. intestine of the' Turnstone . instead of in the bile duct and gall-bladder, and 1yc1cxo Zum iHvnt1 crnu 1ong in the intestine of the Magpie instead -of inI4"the lungs In and air-sacs. each case a reduction of . size was but that P nr as as evident, of prnnonephal urn was not extreme -that Cyclonae The in the latter under- of . changes case are easily cavity stood: Cye1 f in the inhabits a spacious _oelLn air-sacs , with small need for attachment and the ventral sucker and ' oral

sucker are both missing. The body'wall musculature' -is 'reduced as

little movement is required. The region is oxygen-rich and dif- ..reduced muscular activity means that sufficient oxygen, can', , fuse in through the large surface area to satisfy the parasite' ;. ý'maans'that, metabolic needs. Similarly, the reduced activity _Ia the smaller quantity of waste products need diffuse out 'and,.

ratio is adequate.

It is believed that the parasites were coughed up and 'swal- ; lowed. ' The change to a virtually anaerobic environment has

resulted In a smaller body size so that the consequent reduction, -.

in-surface area (which at the lower body volume is 'proportionally

increased) becomes. 'sufficient for gaseous diffusion.. The" change ,. to an active, environment where peristalsis has to'be: resisted,.,

has resulted in reduction of the overall the '. , -a volume" and effec-,,. tiveness of'-the-, body wall musculature in maintaining the trema_, ',

tode', s ±n 'the ' intestine, 'is increased,:; . -position . proportionally °,. the the size is, to Only ovary size and egg, equal -those of trematode_ in, the original location. '.,ý- . __`. -, 551

2). The the ,' parasite mac o.-environment'. ý a)'. The host) s mode of 1 ife. .. ý

If the' host itself is limited to a particular' ecological then the niche, whether migratory-or non-migratory ý parasite', is limited by the host's dietary requirements and by access to a suitable intermediate host. Should the host be marine, ý, living and feeding from the sea, and only touching" dry land for breeding purposes, e. g. the Manx Sh©arwater ' or the Albatross, then any parasite is limited by the necessity of a. fish inter- mediate host, the migratory routes of which coincide with : those, of the hosts and which is -a surface feeder.

The arboreal life of the Paridae similarly imposes a lima=

-tation In that the intermediate host must be an insect. The trematode i. e u_ h1 or idi am has adapted itself to a- terrestrial - host In that the sporocyst is visually attractive and, 'the - . e,_zE z gercarial stage is eliminated.

A habit : of the host such as Anis n1 ntyrhyr_ visiting. >-`... flooded pastures, makes it susceptible to parasites not , nor- orally found-in other Anatidae and in this survey, AnaI 11ýr la P=ronem contnrtn and eras-qum were recorded ' -, -species''using a terrestrial oligochaete as a carrier 'and-intermediate host respectively.,.

Diurnal movements of from feeding. grounds to sea-birds , . roosting areas brings ~ about the -spread of; infection and this` is at particularly ' noticeable : places like Skomer. -Island where many of:. the gulls fly-daily to, Milford, HavenDocks,, feed and ..' on discarded fish from the fishing fleet. " This habit would for, the infections strap t account presence of such' as -ara trldPntnt,, which is otherwise rare in British gulls and: presumably utilises a vertebrate intermediate host since most. ' records are in fish-eaters such as the Guillemots. '

b). Thprnjgrat ions Q the hont

Many Russian workers have shown that migration results in far fauna than had pre- a greater variation of parasitic ., .'ý This is demonstrated 'ý, viously been realised. most clearly -in ; .", the north-south distribution of terrestrial birds-which arrive with parasites of southern origin which are incapable' of infecting nestlings because of the lack of a suitable-

intermediate host. Nestlings and adults become infected by', local the following year, parasites which are missing 'when -the' picked -during birds return. There are also transitory parasites, up.

passage. On the other hand, certain species are- ubiqui-. tous throughout the range and year, such as iia in the Swallow, in this is ui o recorded survey. -... :. 'migratory., ý Sea-birds are less affected by the north-south is by the in, fiat range as demonstrated occurrence of AqQurAt DU in, the' Manx Shearwater in this work. This species has : pre-

viously been,, recorded near Japan in the `Soot"y Shoarwater, which, in New Zealand_ and' nests Southern--South America. - 'The range .'. the, two` birds the,, 'north-. of overlaps -in the Mid-Atlantic; and, =ý. migration south of the sardine intermediate, host and hence the" r'. distribution the parasite is' limited to of . not zoogeographicý1 " regions as it is in, terrestrial birds, fly from the , ý which 553 'Paleartic to Ethiopia, or the Orient and back, each year. Migration in the north temperate or 'Holartic-'is not' so recorded, affected , if the records of rhinos phii a -virguln in Britain Eastern ' Siberia" by'. Belopolskaya,. ' in Turn- and -the , stone are (The reasons for-considering '_ accepted. writer's ". .. Skrj-P_hij=ve_rmj_.; tz Belopolskaya 1953, v ngi _ul a synonym of - E. virtu a are given in Part I,, Section A above). The migration in the reverse direction has brought the first'"

European records of Par rontng_eph_1 urn syct i im. This is

of great interest because, besides demonstrating that parasites;.

are not lost in east-west migrations in the same zoogeographical

region, ringing experiments by ornithologists have ; not-'shown, the Turnstone in Britain beyond" the that wintering migrate 3 ., Ural Mountains. This parasitological evidence, therefore,

increases our- knowledge of the host's habits

c). The nNindancp of the hont.

It is normally stated that the more abundant _-the ; host

the more plentiful the parasitic infections with regard to "._ passerines, both, numbers, and species. Several of the more: rare revealed such as the Raven, have been examined and they-, are to; '

be just as. strongly infected by a number of species and ".local " ._ý dietary' factors are evidently more important.. than the abundance host., Where of-the climatic conditions bring about -a'; massive,,,, -,,,-', migration of, foreign birds into a then the: -. "locality, possibilit infections of alien among local birds` is increased., : Although:,.,. ' the bird itself is to be the interesting .c onsidered. -`a ýmigrant, yigtitarn' in, record of Qnryop .ea a ,Blackbird, in Pembrokeshire 54. in this survey, ' demonstrates that-provided a'suitable inter7,,, host is mediate -present, °an abundant local - species' such as the Blackbird could become infested by unusual an _"parasitic',. species. The parasite's range is extended as a' consequence- of the abundance of this host.

d). Girr; hienl fnetor. ".

The north-south migration of the host over such' barriers'

as the Sahara desert and Atlas mountains which- separates they

Paleartic ' from Ethiopia, or the Himalayas, which separate the Paleartic from the Orient, limits parasite spread in many,, instances. That in the same zone` does, ., an east-west migration -' - not, was discussed for the trematode in the Turnstone. above _-,,_ A geographical barrier such as an ocean is not so effective if the bird remains. in the same zone and this has induced the='f writer to the introduction Leu dy postulate possible of -h]or occurrence spp. , from the Neartic in Section A above. The ' of several of the in this is ý parasitic. species survey related--to:; .,, the mild location the, `, , winter climate and esterly of county, ',--",,, but the cooler, damper limit infections ., summers conversely summer of parasitic species requiring insect intermediate hosts. ----,,, ý' The parasitic fauna is by, also affected . geographical or.,: geological limitations. The trematodes' which% small - numbers 'of utilize marine, and' freshwater molluscs has been -attributed. in:, to lack part a, of lime in local' waters earlier in this part of the work.

57, In r odun ion

To qualify ' for a grant from the British Egg Marketing `' Board, the writer conducted a survey of domestic birds'' in __=.. ý-, Pembrokeshire. In a, county which' is predominantly ' agricul-'_

tural, the, of income derived from, early "_" major, sources are -the potato crop - and , from milk production. While'- many farms main- tain flocks domestic, fowl of varying, there' of sizes, are. _' comparatively few farms of, any substance which concentrate: '-, ý

specifically on egg production or stock rearing. Those that `--, concerned do so. are principally with, the Turkey, (Mal eQgr

pÄ11 öp ý L, ) and the breeding of'. this bird has become 'of,., ,, , significant economic, importance. ' One farm included in' the

survey, maintained' 101000 birds in-1960 and-401000 birds, in_-"- also a of 1964. Such farms may retain shed ýor 'two', fowl, =but rarely more than 500" Only the' sma11' farmer and the owners A, of small-holdings depend much on the domestic hen (Q. 1n. ;. ' eggt i dom_ L. ) of the latter'. in South-Pembrokeshire ýýQ . and some house birds for a, branch of the', firm of Sterlings 'at, -Pentlepoir,

Saunder sfoot...... At the beginning of the, survey,, which aontiüued from July, 1960 to December, 1963, the to trace writer endeavoured , . any important, holminth disease had occurred ;.; outbreak of _which , _ the ' The County Poultry -Officer, --in _County. ý the Divisional :Veterinary - Officer, Mr.; "J. F. Taylor, gave, - '- and .. ,. were helpful advice but both stressed that helminth outbreaks

of indirect concern to animal- health in, -the county-, ' particul-, ":", I-

arly as stock maintenance was, ofa'high standard, and . 558 were regularly by o.farmers. The ant-helmintics administered ; :.. Veterinary Investigation, Officer for South Wales, Mr. D. R. Allen,

provided the following information of individual' post-mortems where significant infestations were diagnosed. The ý number of r =° cases' forthe period, ist October, 1958 to 30th September, 1959 " were Ascariasis 15 Gizzard, worm (geese) rI Tapeworms 11 ý gýmu. trn I _heý .. fl hi An" r1a i l In ýýp r` a3 qD rsar,, The the counties of Monmouthshire Glamorgan, area covered was . Carmarthenshire . and', Pembrokeshire, It was ' evidentthat'- the,

proportion of serious infestations which the writer ' might

record for Pembrokeshire would not, be high.

The writer subsequently, wrote to eleven Veterinary 0ffi's in the to beinformed of any outbreak of -; county asking. rhelmin- ,, _-=, thiasis which might'have arisen in the subsequent, years with 'blackhead', in the-turkey. the end particular, reference to - At : ý= ', of the. period under, survey-no information-had been supplied. manager Finally, the- writer visited the of ' Starlings at provide- ont" Saundersfoot who refused both to information 'of breaks in the birds' 'his care, to the fowl 'to. 'be under arid, allow , ,, included. in the survey.

The results of this survey are compared with those of

previous in the-United Kingdom and two-such surveys _ -workers have been published since 'the completion "of-: the" work - Wakelin

(1964) and Norton-(1964). No' survey of ',this nature has-hitherto';; surveys been conducted in South Wales'*' Most are- the ; result. ',of

laboratory, post-mortems of diseased' or casualty. _birds', sent -to, " centres, the major. except ions, being recognised poultry research , Morgan and Wilson (1938), who give-details of two farms,! A'_- 'B', Owen (1951) included three and and who categories of - management in addition to autopsied' birds. In general, the

previous surveys have related Infestation either'-, to manage-

ment, - Owen (1951) and Wakelin (1964), or to seasonal incidence Morgan and Wilson (1938) and Norton (1964) ., r This is the first -survey, therefore, to be, . major conducted` entirely-on healthy birds ' in their habitat, and to relater

seasonal, variations to a' specific type of management "instead

of to the. ireidence in diseased'. post-mortemed birds. hods

The -survey proceeded- in two distinct periods; July, 1960 ý., to December, 1961 and January, -,1962 to December", 1963. In sources the -first -period faecal samples were collected- from- all to variations but tothe with attention given seasonal not - ,..: type-of In, the 'categoaries ', 'management. second, period selected, _, =_-'of were given more detailed consideration. -management Poultry turkeys were treated separately., This wax ,'_ and _ . due to the great differences in management. -' Whereas the turkey reared is consistently in litter,, the ,poultry are, raised, under and a variety of conditions and with marked', variation in care concern. To -combine both of results'ýwould , sets . -mean'that'_the-, _ differing records for chickens would be made unacceptably, more uniform by the more constant records for the turkey. The', domestic fowl is -dealt : with in 'Section A and the turkey, is ,,- considered-in Section B. Geese, -;ducks and Guinea, Fowl are ale 0',, the together to be found -in county , with varietiesý_such: -as the- The Mandarin Duck. numbers are not sufficient to be ' included' ;,

in a full scale survey and the last-named, birds seem to have, the status, of_domestic pet.

The intestinal worms -recorded, with the over-all total infestations for-turkey fowl during both and periods area ° ., HntPraki agall Innrum (Schrank, 1788) 73.9ý ... " gsenridiaa11I (Schrank, '-1788) , 35.1%

(`, apfII rla`obesignnta Madsen, 1q1+5,, 45 -, _, .O 5% r: f (Holm, 1858). 2.6$ nn111nri, _nudi nta It will` be observed that all are nematodes. by the ' -The 'method: of estimation was examination of 'faecal', ' specimens' were taken as fresh as possible' before being; ý, which _ f dried by the sun or worked int o litter., Because of the vast, _.

numbers, often involving several hundred birds per shad ýý.

was possible only to collect, 2 samples per10, birds _ 'managed'',, _.,,,, . 1+0samples in the first - period. ' From-a unit housing 200, birds, were taken. In each case a fresh intestinal and a `fresh caecal

sample was collected. - In the second period, 2- specimens.. for

every 5 birds were taken giving 80 ,samples ,from' a unit, of; 200'x

birds. -,The, specimens were retained- individually, in pieces, of tin-foil : or less convenientl7, -greased paper and, more -rarely, and in These in preserving tubes. -were stored air-tight jars of; .. ý the Kilner, type'. priorý to examination. For experimental work ý-. the warm, -moist climate in the ears was- found. to, 'facilitate,,: otnbryonation. _ :.. , To determine the effect of, multiple infections, the,:! ma tode ova were-estimated in. numbers per gram'of,, ýfaec, es. method utilised was a modification of. the Stoll egg counting, ' 561 } J: technique suggested to the' writer- by Mr. 'P. Long, of Houghton.

Poultry Research' Station. Only a proportion. of the samples were so treated-and the remainder were- examined by the:,,

Formol-Ether method, of concentration ova and cysts as modified

by Ridley and Hawgood (1956). This is the standard technique , at, the 'London School-of Hygiene' and Tropical 'Medicine. -- - Because

of day to day variability and the fact that most', 'managements once. each could only be visited in three-month period, there seemed in, the . little purpose estimating actual numbers of eggs::, per gram of faeces. By using., a standardised'-pipette so, that,,, ',, -,:

the amount of suspension on the-slide was ` made uniform, ,a =rough -, - .i to the of infestation, opposed guide relative rates as , _simply'. to identification, of appropriate eggs, couldbe' made by counting;,

the numbers of eggs per, slide In the November-December weeks, when large numbers of ' available birds were, killed, intestines and, tracheae,. were `made

to the writer by the bucketful. No attempt to count individual ' ,ý, was 'but there definite ' between-',,, --` --worms made was a correlation, -'---,:, farms. where',, a heavy infestation was predictable from the 'num-` _- bers -of concentrated on the the recovery eggs . slides and actual _, of large numbers of 'worms in' the intestines. Thus a litter

unit where eggs per varied between 8-10, for- a -the slide adult rtd{, ý4 particular , species,, would yield 'upwards of ; 20 AQc gal1i in 'the intestine, while a farm yielding 2-3 eggs per would rarely reveal more than ' 6-10. "worms.. slides adult , Litter samples were, examined according t6 the,, methed, `ýý by Davies (1958) and care. was taken to 'described and Joyner , ý'`ý 562:

follow their example by selecting-litter near ' drinking : Dour. _ý.. content was tains where the- moisture high and also from=drier

areas of the ' unit' and beneath the perches.,,,,

SXTIOH 'A, Tic, rec! 1lta` of . sorvpy, on the hol ri1nth, parncita,

of the domostic' fowl (GnL1lig onllnn'dcM. L. ) in Pombrokeshire.

The total ' infestations -for both thej -first . percentage and second periods of 'survey are included. in Table C 1. The absence ýrnýl (Davaino, 1860) was of ýavn i nýa "i na , which recovered lbq_. Lewis (1930) and Owen (1951), and Of Rnill iet{njnp- by_Owen, (1951) (Mohn, 1858) --. also recovered -, is probably due to the method of survey. The lapse between the collection" of d.. , examination faeces and ' laboratory favoured, the : disintegration ... ,,

of'any gravid proglottids present. The_, sieving of 'mat er ial

through a fine wire mesh in the Formol-'ther technique also barrier, to proglottids: The - of -acts as an effective eggs R. e_ I= in to those- of A. those; st approximate, size glli, -and ., := of D. proelott nn are slightly smaller than the eggs oi', in-any in H. gn1iiri ; neither were evident slide ; prepared ', ,,- the survey. Wakelin (1964) stresses the absence ',of : trematodes

and cestodes,, both of which require'. intermediate hosts, . from,.. birds -intensive They be-, `ý maintained under- conditions. might . expected from range sand, types of management, however: other' , .; Strobila were not recovered, in the Christmas months when whole,,. intestines were available. .= Trematodes , are comparatively rare in fowl,, in .the United' Kingdom -a fact: not ,.indicated- by,, lists" of species, in the,,, -the Adult Domestic Fowl in ;_ TABLE CI Records of Intestinal Helminths in the unit ed. Kingdom. ;. ' ý, . ýý ,

% Ca ipllaria isllaria State o No. % In- Aa oardiacardia Ca ipllaria ß illaria Trichostron Het ri R llietina Dava a Country -Date.Date. Author. ofý - _ health irds Pected i a. ßß 4 aß be nata - 1us gallinarua cesticus' prcPlottina tnu

mixed 78 2. b%' 91. Uv : ,. '2.6% Wales 1930 Lewis - ..; ,. 22.7% 36.4% 85. E ý Scotland 1933 Foggie diseased 22 - - 11.1 e' 40 0.046; ' 14. x Scotland 1938 Morgan & General 113 36.7 8.4 0.01%, 82.3 Wilson Survey diseased ,. 97, ß Farm A 170 - 30.6% . c% t, Hoalthy 7 ` Fay B 285 - 9.8% 9. tß 73. O;o' ý, "_ ý7ý . diseased _ ,' _

Scotland 1939 Morgan& diseased 230, 27-4%'- 53.3 - 86.6% -° -_ 33.5% Wilson 31,1iß 2.6% 3,0 88.8- 21.4% wales 1951 Owen mixed 267 91.0 27. Oq "41.2% Xý, Southern 1961 in. of diseased - - -'_, 23. Endland Agr. Rep 69.2 33.63, 10.kß 45.3p 60.9 0. T - 1964. 'akelin diseased 289 80.8 Oy 9. 21. % 44.9 1.1% 77.0; 2.0% Southern 1964 Torton diseased = 356 1ýÖ. - Fnglard

Wales 1965 triter healthy 96. E 47.2 -' o it -53"t%' 91. & 6,4 standard text-books. ' Morgan & Wilson (1938) tentatively : ý ._... -: identified-a single specimen of Plagior hi sp., Owen (1951).

found two of Br sp. ' in fowland specimens . aý4 an'l adult two N_ot. (Rudolphs, 1809) . ry ndcötyltiP. 3t en1n . sus in: a-fowl `='Wakelin-'.. (1964) but Norton (1964) encountered - none, recovered assorted' give specimens and was unable- to: -any identification because ýý of the of Other than Owen, -the poor state preservation. . -only . record for Wales. is that, 'of Horton-Smith' and-Long_ (1954) _who-;, n-nbii recorded p? agiorehts is Nicoil,, 1909: in the , intestine, . of'a -fowl. . Di cu cinn Azý.. The inteSti a) i. ýPjaaril An idE3nc - of nn1 ` nemit0_o4.

The incidence for 'both the first second isý and,. periods . shown, in Table C 1., Table C' 2 shows the incidence seasonal . and, period (July) 1960 Decembers 1961)ýý variation in =the first -; and Table C3 the incidence in specific managements in the `, - second--period; (January, 1962 December, 1963). - The 'percentages

in Table C 2are rather higher than those of Table C' 3 for

reasons to be considered under c) : 'The effect of the, 'and i seasonal environment' , d) 'The variation' below.,,, -., l: All, percentages' are higher than those of' previous, w,orkers.

shown in Table C1 the exception of (91, '8%). with . which is lower than the figure of Morgan & Wilson (1938) for,.; ' their; Farm, 'A'. (97.1%) These higher, percentages the . reflect influence, propagation 'of ',more_'intensive forms on parasite -the; -, << have more 'constant conditions. of management which . environmental In Czechkoslovakia,,, Bir ova-_Volosinovic ova. ý(1963), found narum; 47. 82.2% of the, fowl infested with H. g ill and '2%' infested . I C-4 C%i co r- - ' rN I-

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41

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q 566: (1960) in U. 8. S. I with A_ ggý li while Denev ." records ,a range', - infestation with A. ', % of -39.6-96.41 galls'and -'only3 2.8-64,8 "- infestation with H rn111 nn rea. the With regard, to H. g 111n ru. , sources of Lewis (193 0) notes birds (91.00 are not disclosed but from the introductory : diseased would appear to derive from both carcases and carcases sent to butchers. The next highest '', that of 'percentage, sources Owen (1951) is derived-from various and reference. to shows that the. highest -his -Table VI (p. ° 119) ', very percentages, differ- 95.9%--and92,9%, are found 'in healthy birds reared under, (Sources 'B"- 'C') ent categories of management. and . . Both , ID ' percentages are higher than those, of khis source which were , diseased birds (91.2%) Thus, for H-' Pal 'a heavier , " -artim, healthy 'as byby, worm burden is supported by- -birds stated: & Wilson (1938).

The' health of the birds also influences the, incidence of'' (1938) found Aana idin gam{. Morgan &Wilson 30,6% of, ythe at healthy birds- were infected, -- Farm' A' and,. only_'27.4% of. the .:; ý. diseased birds infected in their, survey in ' 1939. Owen were _ (1951) that 50.0%, of the. healthy birds 'at Source ' B' were, _found . infested, whereas the diseased and casualtybirds -'autopsied. `,:.. at source ' D' revealed 17.9% infestation. The, to r relation - of disease -infestation -is not so -clear record a for-CIli r. in nb, gnn n. Morgan & Wilson. (1938) higher--, Ga I Ir amongst'',,:.: ý" percentage -infestation of. spp., - diseased birds at Farm B" (69.1%) than amongst, he'althy bird at Farm ' A' (50.0 %) . These, percentages cannot be consideredk °, Äi 5sß

It is larger than that of C. ob gna-ta and could not have

been mistaken had it occurred in the faecal"- specimens. The-

single-occurrence-of C. e udin in the. faeces; and straw

litter of a' group of birds under intensive, management- is even- host more problematic- since an intermediate, ' i's- essential"--, ',

The age of the birds-at was six -the "establishment'concerned to The birds had been raised'- months -one- year. on`grass, -and subsequently, transferred to litter. Infection must have,,, The occurred pr for " to transference. l increasing-scarcity of :

eggs in, the faeces suggested that the infection was dying out

and, the necessity for an -intermediate host eliminated,, the"' Z the unit. This " Possibility: of re-infection within compares .. with Wakelin '(1964) finding"holminths_- in battery -birds. ''-' -

It is concluded from, the -higher incidence of-- all" nematodes

in this survey that previous surveys, utilising-,, post- mortems

birds do not reflect the. percentagesactually, obtaining under,,, = normal-conditions or, even from the" managements from whichthe" :

disea-sed birds have', been taken. Norton (1964). states that:

the, results- obtained by Morkan & Wilson (1938,1939) and healthy diseased. birds Owen(1951) who included both and, in ;. diseased birds do -their surveys, suggest that not ! necessarily.; infections than healthy birds. "ý The " -, carry heavier yr lighter` surveys writer has ` quoted from both extensively above to° show , _; Norton's is incorrect (with., possible exception -that analysis -the Of C_ 'ob-iPrnta) for 'the species, in. this text, and disputes ";

Norton's conclusion, "I It ; follows therefore that the examination, " Vito', of diseased 'can- be expected'-to" give a valid, ' guide the -birds ý -'' 56b

as significant as those- for -H terakis and Agcpridi a, - however, "_

since'they include, C. rannt ; (Schrank, `1788)"and. (- n ft (Moline' 1858). It has been by Moorehouse ' (1944) . established, that C. enudinflAto requires an intermediate host and Wake1in"-

(1964) quotes unpublished evidence that Q- anntis alsö. ý intermediate host. This being then the,; requires an so, -, " relative` percentages on the respective farms surveyed- by

Morgan & Wilson (1938) depend more on the availability of -the "--, intermediate hosts than on the health of the final host

Differences in soil acidity and access to the intermediate-.

hosts must be taken into consideration before these-, percent-

ages carry the validity of those for nematodes with, a"direct', _":.

life cycle. Owen (1951) shows quite clearly a higher percen-

tage infestation with oh ig At from diseased birds ' (Source 'D' 8.8%) (Source 2.5%) - than from healthy birds 'C' _ both sources represent a variety of farms in mid-Walesý,

diseased birds may well be more susceptible to nbg1gn

The higher percentages of the writer reflect the-, increase in infestation birds intensive of under managements' with,,: _-. -, nbnignnta first noted by Wakelin (1964) and furtherý, confirmed-. -by Norton (1964). This _,ý can be attribut6d to the direct,, life. -,1 , cycle and a formerly uncommon nematode now occurs more frequente than Awl? i in the domestic fowl.

The absence of C_ rt i (Schrank, 1788) in -range and - The, writer: l other birds cannot be satisfactorily explained. ý ist of is'-familar with the very character is"ti shapo" - the, "jogg IV. "" a result of experimental! ' work to ' be described in Part :01 -N 4, 0 dPV ON M ap co I W. _0 "

42

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ýd O 5,70 incidence of nematode infestation. -' Morgan & Wilson (1939) infestation varies considerably showed that,: incidenee - of according' to the particular disease-. but- is usually' less: than, '', in healthy birds. .'. b) :T gäf the hast. '. .. .. Owen (1951) in The ' writer has followed considering 'all',

birds than three, months-to be'adult,, but'to determine, of more -FT `, the establishment of an, age immunity, " four age groups-were Table C" 5. -. Since Herrick -utilised as 'shown-in -'the work of ", (1928)1- (1926), Ackert and Herrick Ackert, Porter and Beach (1935) Kerr (1955) onAn`'ýrtdiä g the development and 1' of . ;ý towards 'intestinal nematodes' has an age immunity always -been, - . ;. age `immunity is claimed in H g.q 1 Carl, by Deo assumed. An _ . Srivastava (1962) reinfected chickens aged- '7 and who ,, weeks, 'infection., Fewer- 3 -weeks after th© primary wormswere recovered

from the, - secondary infection, -'Only Wakelin`(1964) quotes unpublished in Cobs. evidence-for an immunity (1938) Morgan & Wilson "seem unable', to interpret their ownresults `ascribe the light infection ( : oni p:: 168 they . 11.1; ) to the diseased birds ' of the. general survey being over six

in with Ackert. and Herrick "(1928) ', ' but in months, -agreement infestation the table` on p. 170 they show. 30.6% with the in 170 healthy young'' cockerells killed for, market. - This.. Suggests the differences in percentage infestation that lower figure. derives from they presence, of disease and. not' from an3r-1,:,

age immunity. _, Their. 1939 paper, gives a higher.. percentage -' this is, in_diseased birds, 27.1+%,', butt still loss _ "-infestation, , _ ;ý 57. ý previous than in the healthy birds of 'the 'ý year. The percen- tage from birds in' which of, 'for'1939 comas , -'the majority- deaths occurred between the ages of 10-16months ' so-that

an age immunity is not indicated. The to first contradictory evidence an age =immunity for and x' gnl 1 innrun is provided by Owen (1951) and is , shown with-similar results for Q. ahsign by Wakelin (1964).

in the Table. Owan racords'. C- obig its in older birds, oInly.

from his Source 'D' (diseased and casualty birds) so', that

age immunity cannot be inferred from his figures. _ Wakelin. does not,. account for the discrepancy between his °ý. but that unpublished. and published, work, suggests the low. percentages, in birds of 4-10 weeks is a consequence of. the.... ý higher for birds of this age for standards of management and .' the shorter time available for contact with parasites, -' This low partially explains the writer's percentages, for' thee-'same- taken age-group but special, care must be in the interpretation, - Of, these figures. The method for the present survey is such that the 'presence of mature females is indicated. --, A. - 11, only - , takes 50 (Ackert, 1937), H 11 m 'take at days , to mature nr; 3': days least 24 days (Clapham, 1933) and C. oh ig ata`. takes 21 that the (Levine,,. 1938). It follows writer's figuresdo not -ý__ indicate the 'actual worm burden for the younger birds: which, will. " include larval stages. The genoral- increase: --numerous, in infestation with age shown, by the writer's figures,, come - ponds increasing of mature worms present and,; "'", ** with, the, number- . ý`°; is similar' tothe results "of Owen and Wakelin., ý, . s ýýý: An interesting, feature the ý of writer's' method'of , survey ., is, that' while the number of infected birds =increases,, the' eggs per remains substantially° the'-- average number of -slide same or ' be in the older. birds. "This' decline in--egg- may' -less . production interpreted by the writer not. to, indicate' -is_ an immunity. to the age immunity, - but to i"either 'an acquired ýor eggs` presence of multiple infections, where fewer fort,; each is the of species bird species produced as -number , per "increases. (See Table C 10 the discussion under, g) : "The, effects and . of-, l multiple 'infections', below)'.

The presence of an age, immunity in domestic fowl, is not ., easily but the contradictory evidence may ,well be due' - solved . , to the conditions" of rearing: ,The -authors, whose, result

indicatethe'lack of an age immunity all drew upon birds living. " the under-"normal field conditions, with" attendant ý",3ý°. hazards -of 'multiple, infections, disease and good,, bad and indifferent The authors work, , management. whose provides evidence age immunity have, for the, -most part,: utilised of an , . There 'birds living under, experimental conditions. -, are, =r"many, papers helminths the `great ", by, workers on poultry. where differ..;, infections epees in percentage" between" laboratory and those'' Clapham '(1932+) under, natural'-conditions are emphasised. ,% found heavy",, that, $ it seems, " impossible to buildup -infestations & Wilson, (1939) conclude' it, 'see'ms innr gym' and Morgan: , that fowls conditions are exposed to, -,,!: ý; evident: under , natural - factor 'or- factors tend to. down'this"partiäl°ý.. some" . which ýbreak ý..ý immunity to heavy' info stations-, which, ",is"-somarked in experi--, mental birds. ' . -, .. -573ý' In these `appear to be statements would one reason for - the supposed 'age immunity' of earlier : workers "and, Kerr (1955) ; , ".. , Under field conditions certain`facbors' must, be present whichrý-, counteract the effect of age experienced under laboratory "--

conditions. Alternately, 'age immunity' may be-an "acquire d

ignmunity", irrelevant to age, such- as was -demonstrated by', , _ successive Shikhabolova '(1959) where : infections with, H. g jIi-.

1lart showed reduction ` in the ' number 'of worms, retarded, a , development Super='infections- ' and, smaller average size. intensified the immune reaction not only to the, second" infection , but ' t*o ', Vatne (1963) has, described the- develop- 'the ° first also. . in to and- went of a strong immunity chickens ,Hg 11 1nirum it appears that the of, Deo 'and Srivastava, ( 1962) 'should'- results . be interpreted as an acquired and' not an, age Immunity,,

These examples of acquired immunity were ,,induceed under Viand may be less marked in field, `. experimental conditions _the, 'e where, be present. A, small acquired,. other factors would -ý immunity is not incompatible with the general, increase in by' Wakelin, Owen And infestation with age ; described- the writer

if it 'serves to, keep the overall infestation- within,, limits

which do'not weaken the bird. ', C): The Rnvirnn f the hrm - -n In the second period of survey'ý(January, 1962,. - December,:

1963), when the habitat was specifically considered, the writer- related recognised seven categories of management which can"be, to those 'of as, in Table ' C3. Two establish-- - previous, workers, for of categories, a), (ii) ý vab) meets ; were selected each - -. 574 fý.. and c); and two and ten establishments for d) and e) res- '..

pectively. Only one small-holding fell into, category a)

( iii) and the owner has since stopped rearing ' fowl-. Reference" to Table`C 3 higher percentages for' most categories' shows = and.. for H ki . 'This' is due to the especially -ter ,, -criteria used in selecting the, units for the second period., ', Managements

whichwere infested" with', all three species were =selected and from these, managements in which the percentages' approximated the first period to 'those for each, nematode in were'-given 'Fstablishments detailed, treatment for, the variation. , -seasonal .. _, different 'percentag'es with only one or two species showed quite ýý and lower than when all. three species. were were usually much . Thus H-terPki present, but with a higher- egg count.. - where infestation was 66.8%9'but alone was present the highest there the ide Cats b)"! - se were-up-to eight ' ova on ' sl s, ,- : mi- intensive. present the highest _ = Where eg, r'tala alone was, Y. ý. was a) permanent" grass' percentages +2.3Aý -y r °= the highest pons, and where only, otg ata occurred, per- centage was 45.8ýý Galup= : a) . straw-1 itter. indicated-on themap: The principle farms visited are, ý.: have (more than type Not infrequently a' farm would one . of in infection management, a factor of- greatr value correlating foods rates under the same conditions, of climate, etc. month's Where this of, up to 3-6 war-a',,h ou sedin " occurred .-birds and later transferred grass pens or 'semiintensive conditions ,` The-' to litter at the 'c'ommencement. of egg-laying. -,. single a farm.. record of C. caudinflata occurred at such 575 Two faecal specimens were collected for every five

birds, 'a caecal and an intestinal. ' In terms of management, the total birds housed the total number of and, number. -of--- specimens taken at random were as follows: - Total no. No. -Of naPP-rl Sa F,

a). Int .,, _naiVA straw. 19000 ; ý400 ( i) -I i Ar - ., (ii). . 7 i. .. non-straw -800 3203$0" iii) -er 200, '.. permanent p_naa es  b). 1ý litter & grass rune 000 " 400 . c) Rr nga grass only-. 900 360 8d)70,348 l-hold nr/C erlfar ä ". e). _Pr v to hnu n -ql coopssemi -"_ permanent runs, or, permanent runs, 150 60

grater number. Had it 'been possible to take a' " of specimens

at more, frequent intervals from all the establishments in each, -, category, ', different might have been" obtained.; While `"";: results , r great no details 1vere'recorded In"the-first period' a" rdeal, 'of.

Variability and for each there were estab:. ' was noted 'category,: -species, infestation for lishment s with `a high level` of. " each, and iii) b)". others with `a low. Categories, a) . "( and c) three, "- always showed infestations with all 'speciesand'., these., ý_".. " medium 'infestation' varied between-a`low and ,aý rate ' of with. an 1. infe"stations, average number of eggs per slide,,, e, chronic at, low '.,, The. intensive. Categorieswith 'litter, a). a -level. "ý{i)"" ii) "showed greater: variability with ` either, a very and much , or#-, ": low infestation 'only aý few "eggs per slide . rate of, and , 57f`. level, others equally well managed, a high of infestation and ._, eggs Categories d) and number of. per slide. . e). ,c onsistently yielded' the lowest percentage but often: with high` egg counts'-, ';

which suggests that individual- variability is more, marked in

managements where' less attention is ýgiven, to the rear'int-Of the fowl.

The trends establishedby Owen, (1951) and Wakelin, (1961f)

are seen-to be borne out "by the, writer results. ',,, Thus

higher level of infestation obtains in the intensive -systems for ignat3 in Category b). A- - gall and C oh especially ý whore run are combined. The 'level of infestation,:, a -litter and , - lower in ii), than in, the, for H. ) 1i ru, m israther= a) .t -ý ies due to the fact that ... other intensive'categor, peat moss, does:, not provide 'ideal conditions, for, embryonation. . d)s in The aP; nnnn inn jjnnc ofInt ctinn nnrrrýtrades

into four sections. and the 'Each. ' year, was =divided 'results March, to July to are, listed: against ' Januäry't0 _April,, -Junes Septembers and October to December. For each three month records for Pembrokeshire, published period, the appropriate- in the Monthly Weather Report of the Afeteriologicäl Office have The temperatures -listed ' ' been obtained. `, are . as maximum, average derived minimum "and difference from the from a thirty- for"1960, 'ands year period: The published results -_1961, 1962; those, for 1963 to 1931-1960. relate to the- period 1921-1950 and` '° form, 'of Because the years 1962 and 1963 part. a=continuous', this text-adapted, the -temperatures survey, the, writer has", ''-in 577

or 1962 to, correspond to, the period '1931-1960. The, tempera- :.

tures for 1960 are, recorded in Fahrenheit and the remainder, are in Centigrade. It has been necessary to, altar the temperature to correspond of 1960 to Centigrade, to . the ,,,' remainder of the survey. '-- Rainfall as ' of inches they difference is, recorded number and . from the average for a thirty-year period. Again the-writer, has for. 1962 to the period `1931-1960 toi, related the. figures , 1963. There correspond with the figures for are fours weather" : - stations 'ins Pembrokeshire, three oný the -West coast'- Brawdy, in-, the, Dale and Milford Haven and one county centre a Haverfordwest., It is considered that Haverfordwest results the, the There are more,, appropriate"for mass of county. are no records for hours of, bright sunlight available'. from the Raverfordwe but this, is not. c onsidered significant -. st station , in way since there Is-no evidence that'direct, sunlight, - any - The ý 'ý, Influences the embryonation of nematode, eggs., general '! `,, stimulus under and is of heat ' is available ,-overcast skies frequently perhaps more constant in litter which receives is no feasible reason to little-, sun. 'Certainly, there suSpec from direct sunlight such as the any advantage obtaining , ,;-ý" sai ati ,, Linnaeus, 'hatchings" for in hý 1758", ` mechanism . >; rainfall The in and the variations, rtemperature _for given in Table C" )+. The, tendency for, cool ' entire period are ., the of the gulf-stream, heavy, thunder-° 'summers under, influence . - months. Is evident from" the storms and gales in the summer . figures.. conditions in Pembrokeshi 7ý TABLE CA summary of weather central from July, 1960 to December, 19639

% Actual Ave Diff, from Actual ins. Diffe from Period Temperature AT, tamp. Rainfall. Ave rainfall-

1960 "+ 0.5 5.07 136 July - Sept. 14.7°c .

1.9 7.12 131 ot. - Deo. ' 7-90C,

1261 ° kar. 7.0 C' 26: 3.59 82 an . -ý

i"0 3.46. 140 Apr, - June 11.9°C +

+ 0.7 4.15 ° July - Sept. 14.9°C ---108 .;

2.4 4.92 94 Oct, - Dec. = 7.4°Cf

1962' _ '3.50 89 Tan.. - liar. 5"06°C _-0.54 , .

° 2.44 98 Apr, June 10.4C 0.8

4.6 July - Sept. 10.6°C -

3.33 63' Oct* Dec. 7.3°C 1.0 jaI6 0.4 2.79 -70' Jan. - gar. 5.2°C

120 0.4 2.96 10060C11, , pr. 'June _

July 1.0 2.72 Sept. 14.2o-C', - -,-71 -- -' 0.1- 4.87 93 Oot. -Deo. 8.4°C + 579 (j), First P rM (July, 1960 - December, 1961)

Because of-, the overlap of years) it has. been necessary temperature to average the and, rainfall figures shown -for July-September, and October-December 'in Table C' 4 Table, C . and 2. The temperature for July-September , was above average in both . years while those-'for October-December were less. in, each year*, -,,'

December, 1961 was the coldest Christmas of th'ecentury, ýand took the average for both years well be1ow, 'normal. The , tempera- tare-for October was the thirty-year_ for above average seven .°, years in succession, 1957-1963. The rainfall--for , the July-. °

September period 'was above average yin both years but November and December, 1961 ' were , below normal.:

The total, infestation for, respective species -in, first ý,. -the period is higher than those-- for the second period.. (Compar©,: This is, due'partly,, to the of Tables C2 and 3) . effect ' the_r". average temperatures 'the" ý above and -partly -to effect ,:of com-r. biasing the various systems of management in the same results ', for the-first period so that so rg variation, is Thee<' _ , eliminated. -.'. ' number of per=, is also, higher, than, in the" average . eggs slide second- period. The incidence for the , by Morgan various - species recorded & Wilson (1938)., and Norton (1964)' are compared- with, those of the . ý writer, in Table 'Cý6. , Th©re`, is -a report of seasonal variation for if. gall inarum in, domestic fowl'. in Russia (Savchenko, 1959)

A record was kept of-the incidence -in, ten hens. '.,,,The monthly , lowest figure occurred in January-Parch, ', then there was a the gradual' rise in level to July a August after which :`,, 86, aö _ äo 1 4 co

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CA 58 percentage burden gradually fell until December. This_ is the

accepted pattern in traditional methods, of management. Birova-_

Volosinovicova (1963), however, records, the highest incidence

in, winter and spring months, a trend also-observed by Norton

(1964) for AA_gni1i. Morgan &, Wilson-(1938)'ýand- the, writer . both found an equally high level' in the October-December', periods - , ý , and the, percentages, for January-March; in this : survey were not '.: lower than those ' months. much of -the summer The higher- figures 'in this for, the 'cooler: of survey, - months . ,' the year can be attributed to the exceptionally high rate'"-ofý, warm, embryonation -occurring in, the wet summer. This would

give, a strong incidence of secondary infections, in the 'warm September, October As and - 'months which äevide ntly maintained. through the in the birds. The ' fowl in litter benefit from winter . : external warmth' and litter conditions are '-such that further ý- -this _ infections the small can occur, in winter months _when', the climatic conditions lower the 'incidence in"-birds under, non- intensive , systems of management. ' Asa result thaverage percenta'ges- for , ,. January-March is higher than previously, recorded in the : United,, ý

Kingdom. In the second -period, this is more clearly. demonstrated.

Norton (1964) the highest in'' records -incidence' A11 - in the January-March October-December Morgan and . months, while ü,- & Wilson (1938) state, 'no marked seasonal' variation. ' A small J': - . -: ý variation' is indicated by- the writer the ;.,. seasonal -but again conditions, obtaining, in straw litters raise : the inn idenee in -. colder months' above. that which ' might:, be expected. Birova-', ", Volosinovicova (1963). -like found the highest infec- -the', writer-, the ,t ions in late spring. 582 The_ same pattern of infestation, is recorded by the writer, for-, fl Morgan & Wilson (1938) that the ahoi ýt . state .. highest period of infestation , isin the April-June months with

a, gradual decrease towards the' early months of the --following they, an year. . In October,, recorded, exceptionallyhigh, percen- -.. tage infection.

This in infection rate in October is a_,, sudden-increase , It is, ' 'Norton consequence secondary Infections., ' seen in .. _of _ figures fall below those for July-September which . ofApril-June in the only to. rise again in-October-December, arid . work of Birova- Volosinovicova in Czcchoslovakia, ' 'who, noted two-peaks of,,

infection per ' year in 1b y and, in September. Er After awars summer

_like that" of 1961,. this, is very, evident_ and. is seen in the despite intensive, systems in the second period: of survey - the c old , wet summers of : 1962 and 1963. The high rates of infection recorded'by"Norton for A. g1 inTthe. period, October-11arch;

(1964) reflect the=trend iný_the, 'intensive" systems where the. at worm burden is= maintained a high level, throughout the winter. fit) se nnrl Period (January, 1962-December, 1963). -The incidence for- the various categories `percentage , of. th© two year management is shown ' in Table C 3. During' `survey the the average or, the thirty-year, -temperature was above periods 1931-1960,.. for six, only. Those were Januäry,.,.. _©onths 1962 (O 6cc' October, (0.19C above), February, 1962 , above) , `'196j -- (a. Oct 1963 (0. Vt. 2oC `above), Junos 1963 (0,5oC above) obar ý = . , . and' November, (1.50C-'above) .° While the temperatures:, above) -1963 - , be-thought, relevant ' to -ýý for January and February, 1962 cannot' ,, 583

in the non-intensive"- systems, the any seasonal' variation ., would warmer June, October and November months appear- tv

assist in. promoting primary and secondary infections, in the,

respective years. The warm, temperatures of January and

February, were counteracted ' by heavy. falls of snow and sleet . days January February, 1963 the, on several while . and were -ý coldest of the century other, than 19+7. Birova-Volosinovicova'

(1963) demonstrated the' ability of mäatode, ova to survive:,

under snow. ý... considerably Textreff The, rainfall varied, the s being 'ý 1963 January, 1963 (18.0%) and larch, (183-0%). ' The', months:

April-September' 1962 had, above 'average rainfall =and then`. a ,,ý, '' -:,_. relatively dry period follo d from October, 1962 (39. ) to . a period February, 1963 C55" ). July-Octobers 1963 was also ,:

of below, 'average rainfall. ' Except in drought, in' ; able - whether rainfall is as significant. embryonation as.. there isalways rainfall temperature. - Even in hot rmonthsl some and often' exce ssive ' dew. , Inevitably therey is moisturo ýin : the': ý. , even cif-, micro-climate ar ound -the drinking vessels the, remainder , off 'a yard or, run is dry. The greater importance of temperature,

in-hastening embryonation is shown by the'- work of Osipov, (1958). humidity'., for the months. of- }ay ',and 'July, when' general" was -the": ' -The the' rate _of. embryonation, same.. experiment concerned. of, .. ý' ; 'Hg l1 ý rnm ova on the ground in central U, S. S. R. Fm ryon MMn lg t_1 on 31 June 21 ? C 60%', 1 -24 days' 10% 26 days "' July, 15.. C- 2(3- - . August 14.8 C 81% 23-30 days"' m

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le-ýb L r i " 2aä P 4O q , y A cal x r-1 yý r-1 ,. 4 2 ýiF -; 585 n Because of the consecutive colder summers of 1962 and

1963, 'it is regretted that the work could not be extended, ,. , -. their; the to include, 19641, when effect on incidence of, nema-; , todos in the fowl, maintained in the non-intensive unit sý could

have been studied. The incidence is 'in: Table C 7. All of >Heterakis shown managements showed two peaks of infestation;,,, one in the spring

second in the autumnwith a drop, the " later summer and ,a . -during months. The intensive categories showedý_less variation than

the nor the variation being'-, in (fl -intensive, seasonal ,a)..;, -, 5.6%, lii) 4.8%; Cataonry i1,,, ii)ß - _(, showed greater y _-varia' tion, 7. A as can be expected in the', greater 'susceptibility.

to climatic changes, in permanent grass runs. Of the. 'free- , ,'. birds, c.).. only 6.3% but LI).. showed 22.2% )... range showed and , = 27.7% variation respectively. The influence of climate is -- clear and must , be_attributed, tothe effect of temperature rate.. Thus; the; cool, 'wet, - summers of July-September,

1961, and 1962 depressed the embryonation for the following

year in, the' non intensive systems, but the warmer "Octobers increased. tho during ý slightly ©mbryonation rate - -the, winter', -, '; =.; months. {. Both units a), Lill and £1. possessed coops with These perches. -,, were. cleanod'. regularly but evidently the intensity of infection was such; that. sufficient Ova could

embryonate', in the warmer confines despite' external'conditiona. more' The', influence of - the, cooler summers- is; clearly revealed by the variations in the numbers slide. Thus in of eggs per . (ii 1) and the non-'intensive units, the second peak shoved'-'_ ., _a. . 0 ö s 4.3 ýt ý. 58G 4 14 u1 co L1 & -t * 1 1 1, 1 1 I a0 b ýJ 4ý M 4 - 4fl

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82

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0' m m - t-3 a V1 U of 1 1_ .1 ý'ý ý' " 40 though the infesta- a much smaller egg count even percentage ,: tion. of the fowl rose.;

The percentfge infestation with, Asc zir idii is shown in--- Table C'8. Categories hl,. the and , showed usual twice yearly peaks, with a seasonal variation of 1.8%1-- 7.3% The remaining `;, 2-516-and respectively. -categories showed, a depression after the summer months which the percentage -from increased I the This with-, gradually ýuntil -spring. accords '. Morgan & Wilson (1938) and Birova-Volosinovic, ova- (1963) for, g- in and Bir (1963) for-" A- n 11 gn1 ruin ova-Volosinovic ova - .` It is evident from the writer's results that, '-in the non--

intensive managements, A 11i is more, sensitive' to climatic innrinm. conditions than is H- gill This can be attributed to'. embryonation rate, Birova-Volosinovicova having demonstrated -the, _, 15oC, H- i days -that at gall narºsm -embryonatosin 72.75 while 1 e_a1li takes up t o 100 days. ' The cool, will'-' ,. -, swot, summers _. = down : the "An Idi 'that see slow ambryonation rate of. -ter tn' so :a ond peak is not evident in the non-intensive units, the - and' warm,, October months instead contribute' to 'a slow build up'.,Iininfes- tation until, the 'following spring. The seasonal for variation ,,-: a1., Categories aL ' and were 14.1%, l+. 3%, 18.4$' ., ,. and 10.3%' respectively. , In-. the intensive units, the average number of f ag s per . . identical-with! those of the-infestation ýý- slide showed, peaks - . ... Ina) t) intensive the rates. -t1i and the nom. systems, average :P low number of, eggs remained at level, even when the, actual . .a low infestation rate increased. Thus the-worm. burden remained,, on embryonation. ý-r due ' to the influence of cool, summers .__ 4 588 s.i. Ul % co %D ^' ä ot1 1 1 1 1 ` M ä O\ m to m 1m fl m, 43 U' O M u'1 co m -t , %0 %D UN

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o ci H 40 03 9 p L:

119 , ' ýý 43ý t) IVA. i 1 1 1 14 9 42 4-1 ä : 0 ý, 0 . The seasonal, variation of C pf11. rin ohnignhtn (T ble C 9) t to that A. 'with twice is similar ý of Call a, yearly peak in - the intensive units acid a single peak, infestation in, the

spring, in the non-intensive managements. The least variation is in, (1) the in 18.7%. ý seen a} .` ý_-3.7%j and" greatest " The the demonstrated ' egg counts also followed pattern , for_ A. gnll to

The högeni i int amitodAs. e) t. n . .yof _cti a1 From the. farms in both, the first visited-, and . second periods; ý, information was, requested' of 'any birds dying under conditions , which indicated that death was, due to heavy holminthicý infec-,; ý_,4 . tions. No cases f occurred although'there -weredeathsfrom, owl pest,, birds becoming 'egg-bound' and- other, miscellaneous causes. '

The.. pathogenicity: of intestinal nematodes, even in heavy - infections, is considered-to be doubtful- by the writer. This contrasts strongly with the: Report of- Poultry ', - often quoted of Technical Committee. for Great 'Britain (1938)° which- stated: that

worm, infestation is, one of-the three most frequently diagnosed,,, - causes, of -few death is -death. -'There are records where attribu- ted solely, to--the nematodes occurring-in this survey, under .'; natural-. conditions. ' In elucidating, the survey life cycle "of

C. obalgria tag Levine (1938) was: able to cause ' death by 'massive

experimental infections but in the field various 'factors work

against such super-infections.

The `only pathogenic 'nematode of poultry is', ngnmes trieheý `. -, -'Sy (Montagu, X1811) the ' absence, of which in-the report 'of Owen--1 (1951)'- 'speciesý- and in '-the `present ' survey, indicates - that this 59U. is little iný South`, Wales. ' case of, significance - "The single , in South Wales quoted in Dr. Allen's letter in -the' introduction ,

above was not stated as the- cause of death.., In: the "Christmas the tracheae large, months, ' when of numbers' of birds , were

available to the writenot, a single 'infection was. recovered , from any system' of management. ' ''As mentioned in Part Ij", 'Section" , . C7. it. was not recovered from arty of, the migratory starlings and

it-is a parasite of minimal importance in the : local Corvidae. ' ,'. f); The thA' tnhli_shmnt infaet Otis, ' _nanditiaria'or e of ehrnnla In fatal the intensive Sys-"' . _the absence. of any 'infections, tems, 'appear ideal for. even .,on' well managed establishments, 'the. persistence. of, an, infection at a high level , of' intensity and the , devoid of soasonal variation, more, constant percentages . much is found' in these units. (See Tables C 7, ', 8,9) This , especially not, true of ; the nematodes which do require an=inter, - .., mediate 'host and, reference to, Table C2 : for the adult: domestic -fowl bears out the "statements, of- Wakelin (1964)'ý'concerning the, increase in, infestation rates these , of nematodes, : especially, -= for. G nta. ý: Chronic infections of either a high or-;. low= intensity, both show two seasonal peaks : of infe"station- in' the Infections, intensive units and egg counts are uniformly high. ',% ', peat are less persistent in moss and bracken. litters. than, straw. Dampness through leakage; of the roof, or, ofthe,. drinking: appliances is a major' factor in=maintaining a- chronic'' infection Similarly, the as are infrequent changes of : litter. rearing: of not cleaned least every chicks in units which are. at twenty =''.; 596 ý. days,, tends to increase the worm burden- and it is essential _ to rear chicks on ground which' has not been- utilised for. adult ;, fowl. Birds reared in, pens on the, range must be moved to -clean every fifteen days. ground ,: g): The, effect of mti tja le infecti_on, 4.

A, correlationEbetween number of species present andthe ry`demon- number of. individual, worms in an infected bird has,. been

strated by Morgan & Wilson (1939), Owon (1951) and: Wakelin , , 1964)'- amongst., others. In the_'present surveys, where eggs and'. have not 'the numbers of -worms been counted, no '- actual _ such,, -, relationship is evident. The, writer suggests-. that; the results,; -

of-, previous workers may. not-have the significance, attributed large larval will -'. to them : since, -.-inevitably, numbers of ° stages ', in While heavy infestation 'par- be included any, c ount s. `,- ;a of' a , t the to further icular species mayx.'increase, susceptibility . infectionsby the, same', and, other species, . the success of such-, ";:,

infections. depends =on =the maturation- andproduction of eggs-, by,:, ' fertilised -Morgan &, Wilson (1939) show that for -females. tern Is gni1 iniri1m, ' 'a worm burden does not 'heavy __--increase -.ý.., their -pathogenicity'in healthy- birds and-'before,, the effect-of ý. a multiple infection-can be thought significant : all-larval, stages mustreach', maturity* The `hatching mechanism for, the ova. is unlikely to influenced by the >presence of; 'other - _be, , -, species, but, age "immunity if present,,, greater, resistance the 'heavy' ; breeds, the -presence - of, established , and ýpreviously` . chances- being,, attained adult worms, -must minimise '.the - of maturity, ý, by successive of the same and ', other species.: -, " -infections ,, , The writer has, in fact, recovered fewer, numbers of eggs-:

for each species per. -gram of faeces, where two or, more were,

present than where single, infestations were indicated by the"..

presence of only one type' of egg (Table. C 10)., This, suggests, ";., that the which the impressive many, of stages make up . numbers' 4, shown by Owen (1951) in his. table , are' larvae which do : not,, ' .:

mature because of prevailing competition., If maturity `is not

attained by the majority of worms prosent in any count,. then possibility of such statements as 'the fowls bee owing heavily,

infected' could- be substantially- reduced by,, preventing, the num-' berof helminth to infest ' the birds from species available -". two' (Owen,, '1951), have, little 'significance, the rising above - -in V essentially non-pathogenic nematodes, under consideration. ° The _''

worm burden in terms'-of numbers recorded , iss, in_ the absence of endanger disease, rarely sufficient to the health of the bird'° - V , and, Savenchenko (1962) has shown that super- infection'-leads' to, -,,-'

spontaneous expulsion of $ erik gni 1 Innrum in the,, faeces.

The decrease inthe, number of eggs per gram of, faeces is in part -explained by Shikhabolova'saccöunt (1959) of ac quired'.: immunity to both super-infection.. The effect of the . normal and density of, infect ions' has : been shown by: Savenehenko. ", , --single also (1962) who-fed controlled numbers- of between 5V and, 1ý 000 ova of 11 to An increase in density led to a gg? WE" M -fowl. , r reduc- tion'of length, a reduction -in -length of life and. -a lengthening., of the, time of development. H. 'g h1im remainsVin the host". _ for "a year (Osipov, 1957) is to', be expected that the and, -it . t.-, introduction to the infection would of-further' species . primary , F 594 result in suppression through the combined effects of density and acquired immunity. Hence the fewer numbers of egg per gram faeces per species recorded by the writer.

It is concluded that an increase in the number of species present means that fewer of each will mature and the chances of dissemination will be mitigated' not increased, by multiple infections. The writer's counter-proposition that multiple infections are self-limiting can only be confirmed by further experimental work in which factors other than the number of larvae and adults are recorded.

-;;LTION B. The rg i1 s of survey on the helminth n-Iracitea

the domestic turkey (Iles 11a dom. L. ) in of ri ß;;, .quo jPmbr ök .

The total percentage infestations for both first and second periods are shown in Table C 11. In the first period, the distribution, seasonal variation and age of the host was con- sidered and in the second, selected types of management. Table

C 11 also gives the results of Owen (1951), the only earlier worker to study the parasites of the turkey in the United King- dom. Unlike Owen, no cestodes were recovered, even in the

Christmas months when many intestines were made available for dissection. Neither were trematodes recovoru: d although there are several reports from the turkey. Rees (1952) described Sn1 igl Rees, 1952, from E1 ngiorchi s( . afdu1 n_ri__) mara1 o rh 4

Wales, Varicak and Erlich (1941+) recorded g-rhinostom r voln . nm

(Fröhlich, 1802) from turkeys in Germany, Birova and Busa (1957)

59G

reported Proctthngnn nunantüs , (Rudolphs, 1809) An the oviducts of turkeys-in Czechoslovakia', Valente (1958) described rAenrvatern FEhs i nnp rv; hinm (Von Linstow, in' Italy, -, an1=: _1873)_ -'; Paudere (1958) 'found four species. of trematode, ' two species of

costode ' and six_ species of nematode°inLatvia.

The most common nematode in the- turkey is ' et rgi n- the,, in Czechoslovakia, .ar1lý and of seventeen' species occurring =aý Birova-Volosinovicova (1963) lists H. gam = ilm ßP^11-1 ,

C. hnrsýit-i' (considered, enudinfla . a, -C_ a synonym of, ý, =+udinflata by most - r' ob " the common., The: ', authorities) and, -Signnt, i as most . -' least common'were Thn 'r,hn i uni (isnmi1n'. jQy nY --, n Tr iehos, mnPV to 1y i 'grid Thomin-r 'c nl l (Q_ anatia of most _q workers),,. " Vasilev (1962) listed thirteen species ' of "nematodes from turkeys in Bulgaria.

In interesting, the of an . paper, comparing nematode ""burden , --l wild, "pen-reared wild and domesticated -.turkeys, Maxfield, _'Reid,..; "` & Hayes' (1963) found Äscnridi , di nsimtlis Vigueras, 1931=to be ',,-,. turkeys, in . most common in wild i --1 pen-reared wild:.: turkeys H- li domesticated birds.: ` d_ and ý -ý "in dtsniml its =nematode``of turkeys the common' ascarid -,although the only report for Britain is by Horton-Smith 'and' Long (1957) , The size of the egg does not differ from that* of g= "i and males-which could` be', Identi despite tho -presence of undoubtedly, fled. as dj4imiti_'in, the Christmas `post-mortemsthe predominant; species in, this survey Is considered to"be galls. Kerr "(19579 '1958), has-worked on both: species and has shown that while turkeys are more, readily, infected with dis ni mi IQ"; they have a'" 597 j than-is by for greater affinity with shown chickens , sI ; st.

There, no cases of. syngamiasis during - were ý_the, survey and , the nematode was not reported by -Owen (1951), Long (1957) -'des-, - cribed 'a serious outbreak in 2,000 -turkeys ho believed: -- .which had been introduced by housed starlings. :'The', poults , were -in pens and the earth and litter beneath contained, infected housed-",, "ign nin fam. Most of , thee birds in this survey, were ' --, in litter with" concrete or', wooden floors and when an. systems -, , usually -earth floor was' utilized' the layer of litter ' prevented

access by, the poults, to any fauna present. Small, birds such as the' the ubiquitous and-indigenous sparrow -able to_enter ,_are but largerbirds, the the units - such as-. migratory starling and corvids, were prevented from entering by, wire-netting. Thus,

careful -management ensures against -tapeworm infections. '`, Din -uasi n

The, onern1 tneidere of intnstirn1 ne todns,

of the" two surveys' included in Table; C:. 11' -A comparison indicates: that H. ''gallinarum is not as-common in- the-, highly: "- now being practised, but, the- efficient litter,. techniques , uniform. `r I conditions make for higher 'infestations -with', A- l? : than, formerly occurred. The -fact' that A. i has: not the ° affinity= for. turkeys- that itý has for chickens , seen in the comparatively _is-. The, of -low rate of. infestation. careful rearing -turkey yoults has-not eliminated these nomatodes which must, be introduced, from infected fowl at, Owen did not an_early age, --. , -recover, infestation, " C. obs fills. but 'found a relatively high percentage -,,, . . 59S',, 5-II

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-'"-: __ ý -. 599 C- This last intermediate of -a»dinfi ata. species requires , an of in host and the great efficiency the turkey -industry,, - Pem--,, brokeshire is indicated by its almost complete elimination. - _, .,, Trstron y iq t, uI g is similarly of little importance.:

In the first period, July, 1960-December, 1961, nineteen together about ý. establishments were visited which maintained ý' , . 38,000 birds. Two faecal samples were taken for every ten and birds, an intestinal and a caecal, between July, 1960

December, 1960. In the following year a typical unit at each-,:,-: ' farm was sampled representing about 10,000 birds. The percen- tage infestation is shown in Table C 12. In the January, 1962-December, 1963, eight i-. second period, _ . typical units were examined and two faecal specimens were taken,,: -, (Table The for every five birds. C 13). units were selected, ,.., x because they form of management because, they P- represented a and _. showed an average rate of infestation by all 'three of, the more common nematodes. The percentage infestation was therefore,.,,. ;." higher than in the first period which had included a number,, of '.;. establishments with sparse infestations. The total percentages--,, - wore: I- : -s H- gl1 Innrum - 7+%, A. Call 28%, and C.. ob Ig ýtý 45ý. The is in --- number of birds sampled at each unit given -, °- --_ Table C ],3. b) The tja_ . age of hont. None of the establishments maintain their flocks', for -more .'. than ten months and it is not possible to study thet-effect of the age of the host on_the burden. There is, no 'reduction--, 'ti,-, worm 1 of is demonstrated nematode parasites up to the age, of one year --as the increase in in Table Cr 12., by, progressive percentages _- Th S a1 Intn, c). _nao -of, ctinnI Mmntödes, For similar reasons, there, is scarcely any-seasonal varia-, tion. 'All species show a peak infection in the April-June months and there is a slight host reaction evident in, July and August. The intensive conditions favour repeated re- and infection and an increasing number of ova are passed poults__, only, infected until the birds are killed (Table C 12). The' exception is C. eiudinflntn which dies out owing to the absence of the intermediate host. d). The the hont. environment of . -" (1955) that the in the Davies & Joyner found increase ,'' moisture content of the litter in winter months caused a great increase in the idial oocyst from November. to c occ population ,` February, and the progressive build-up of nematode infections: _',°. in be due to increase in humidity, is seen which must part an <<: v_ in Table C 12. The drainage of moisture from litter is 'impor- ,. tant and in Table C 13, it is demonstrated that the establishment floors lower infestation with earth support a of nematodes -than'-, ýprovide those with concrete floors. (Units 4,5 and 6). Straw-litters `.

the best medium for embryonation (units 1,6' and 7) ý with peat moss as the least suitable medium (units 2, and, 8) ý, An important factor is heat absorption by the environment_ in '-_:, summery and retention in winter. The prefabricated buildings are all constructed of a non-inflammable concrete/asbestos commonly The converted barns-. both have tiled':,:, mixture utilized. , 6 01 TABLE C 13: The % infestation of: the, turkey with intestinal nexiatoüos (Januaryy1962 ý'--- related to different oatogori©e of: manaGem©nt. -'' December 1963 ). ý ,. _:

No. of Hetorakis Ascaridia Ca pill pia Type of Management. Birds, rall- inarum ells obs rpmt,a

(1) Prefabricated building 67 69, Concrete floor - straw 500 92 litter. :_'

(2) Prefabricated building

Concrete floor - peat 500 73 18 35 taoss.

(3) Prefabricated building Concrete floor 500 87 43 ýa63 - sawdust . and chaff.

4) Converted barn ;' . Earth floor - str= 250 75 30 51 ý -°

(5) Converted barn

Earth floor - peat 150 46 11 20, '"= _ Moss. .

6) Lean-to outhouse Earth floor bracken 300 81 37 ' 58 - .. and straw.

7) Corrugated iron unit Concrete floor - straw, 300 74+ 23 ' '41 chaff and sawdust.

8) Corru ated iron unit Conorete'floor -peat 500 6ý ' 15 ' 23 moss with somo earth. 62 roofs and brick wa11s, while units 61 7 and'8 all had, corruga-, ted iron roofs. Units 2 and, 3 are cooler in summer but warm, -12 in winter when heat and moisture together, are most important'--, for embryonation. Similarly the tiled roofs are cooler ný. a summer but warmer in winter. The units with corrugated iron roofs are very hot in summer and dry out the litter and area--, cooler in winter through heat conductivity. Thus an ideal', ' unit would permit absoption of moisture via an earth floor, brick walls and a roof which permits some conduction of heat, ' with peat moss as the principle component of the litter.

However, there is generally more leakage from corrugated= iron roofs than with tiles or a prefabricated roof. Thus unit, unit showed the lowest infection rate. Least suitable is 1 in which a very warm humid atmosphere is maintained. These'"'- conclusions assume that there is little leakage from drinking-_ appliances, but several units visited possessed open 'drinking, troughs and litter in the vicinity was very wet. Ruf, (l95): ; " has shown that Hat-ras eggs can only be dessicated''at' a' sur--``. face humidity of less than 5p. A- g nI is more. apparently, .ý to humidity than Her IcJ Cipill In bRae, r susceptible or ."and= . (1935) stated that a humidity of 82-86% is required, for the complete development of the ova at 2200. The only lethal factor for ova in the litter is "dryness. ý ,. Thus a high temperature and lack of moisture, must bethe, , cheat of the farmer. Chronic infestations are propagated whore `. and a high' humidity are found together as in unit, 1:;, e). ý athogenieitrr of fntýstiný name o a.

the, death- be At none of establishments could,, , attributed" of solely'to a nematode infection. On one'farm a-, shed -500"birds, died from blackheads Various workers have shown` that

naked protozoon is not infective and that it must be transmitted '_ (Wehr, 1954). In the high rate of- through Hn Arnkig ova view of . infestation with Het rakis, enterohepatitis is rare in the $eteraki ',, ý county. It is known that only one in 200 ova ,'will contain the protozoan and at present an even lower ration must

occur in the county. The migratory larvae of IL Cal-1-1 are

destructive and cause emaciation and death. The relatively to they ` low percentages of this nematode can be ascribed small"', -f the turkey deaths due-,,, affinity with and mitigate against 4°r this species. Egg counts for this species were below, those 'f orl, C Except in the He ._ nkis and pii1nrii. case of super-infections, to ý ÄtÄ. artificially induced, death is unknown with regard Q.. oh

rnrn,

1) A higher percentage infestation of intestinal nematodes in the domestic fowl has been recorded than hitherto. This ' is

considered to be more representative of the worm burden

healthy poultry in their environment than the results: of , previous

workers who have recorded the percentage infestation in_

and diseased birds sent to centres for post-mortem.

2) In agreement with Owen (1951) and Wakelin (196, the present;

survey records no reduction in infestation with the increasing

of the host. Any numbers occurring is "con-' age reduction-in ; to infections'and sidered be an acquiraed immunity frön-previous not° the increase in the ' " of the host '_ý :_ -result of an ý- age . ,:. The influence of is -towary, considerably. 3) . environment seen depending the -birds : maintained in intensive or .ý"ý on whether are , . non-intensive conditions. -Intensive conditions -favour repeated ,:"-re- ý. infections' regardless of climatic conditions whereas non-- a intensive managements maintain a lower infection rate-' and show, -''

greater variability in intensity.

4). The fowl in intensive conditions show two peaks of infec-,

tion year, one in the second in the early.., per spring and a , .` suppres- autumn. Ascnr tdil gaIll and Cp llarI obsi grritg may be

sod altogether if a cold, wet summer depresses the rate, of embryonation unduly. Correlated with this depression, isaý

decrease in the number of eggs passed. 5) There is the the intestinal . no evidence of pathogenicity of nematodes recorded in the survey and y n: ns trnnhen did,, not

occur.

6). It is considered that the effect of the multiplication of .,

the nematode species present is to decrease the number, "of adult in based the "of ,; worms of each species, evidence on number -,ova ,H for each species passed in the faeces.

7). Turkey poults are less strongly infested with intestinal-;, "', reared nematodes and the uniform environment in which they are

precludes any significant seasonal variation. 8). Chronic infections found in intensive 'main- are units .which tain high temperature with high humidity not---,, a coupled a and,, are , ' influenced by climatic variation. Regular forking of'litter`; ö, disperse the the litter will heat-of fermentation and cause ,, dry out, thus slowing the rate of embryonatien..

Tnt oduetion

To the of the British. Egg ""'. comply with -requirements . _. Marketing Board, it was necessary to devote time to experimental.

work with poultry. Dr. C. Horton-Smith of Houghton Poultry, '

Research Station in Huntingdon was appointed supervisor, and-:. ''

in discussion it was decided to link the taxonomic section of

the thesis farms described in Part II and with a survey of as r,_ to an estimation of the importance of wild birds in the i dissemi-.. = nation of helminths of poultry.

Apart from C. o sig ata Madsen, 1945, which has been from Passeriformes, Galliformes recorded a wide variety of and .;. Anseriformes, there are few records in the literature of het-:. "_` ' minths which occur naturally in both wild and domesticated birds. °,

Gn1l rrinh rn Cahn Ward, 1917, is found in subcutaneous cysts in -

the house sparrow, starling and other passerines and it has 'been-: turkey North recorded from the fowl and the in both Europe and', , America. Several Typh1nnr 1 irn spp. have been found in waterfowl'

and in ducks and geese. Echinostoma revolutum (Fröhlich,,. 1802) _-, inhabits the chicken and the turkey in addition to many. wild

bird species and there are records of HypUderAen onnide im in the There (Bloch, 1782) chicken. are no records of , costodes-ý,, _ - infesting both wild and domesticated birds and very few referring infection. Miller, ((1937)' to nematodes except by experimental -

attempted, unsuccessfully, infected pigeons with Asearidfn (Schrank, 1788) H i (Schrank, 1788), and and .t . rnki a ßn11 nr im ' C Stoican (1960) infected chickens with pil . riý enrvorýýn - Schwabe (1951) was, able to infec_ t (Rudolphi, -1819) from rooks., - ,, . _, (Q imb five species of wild birds including the, Rock -Dove 1! 1 iy a L. ) with, Oxy-g irnra mar o310, decided that wild'birds -but are of little importance in dissemination despite its natural occurrence in five local species. Gibson and Barnes (195? );: `. in domestic ducks found Aminrin urn -i n geese and and con- that the outbreak originated through contact with-wild, sidered , waterfowl. Numerous workers have shown that it is possible to infect poultry experimentally with Syngr rnn treachea (Montagu,

1811) obtained from rooks, starlings and blackbirds (e. g. Taylor,, -, _: 1928; Clapham, 1935) "= 14i t M, hods _rinl n and . Two lines of research were decided upon with Dr. Horton-,.. ý.

Smith. Firsts the discovery and extent of natural infectionsby examination of wild birds. This was covered by the dissections, for the taxonomic work but, in addition, dirt from claws-and,. s., . beak was collected and examined for nematode ova. Particular'-I, attention was given to those passerine birds which are naturally. associated with range birds and poultry runs; rooks, crows ý;'ý; jackdaws, magpies, sparrows, chaffinches, blackbirds, starlings These and, less frequently, thrushes and wag-tails. birds 'can:.. ,, be divided into two those which have extended' flight groups: an _- range over five square miles - rooks, jackdaws, crows and and, starlings - and those which are localised at one farm : while ýý= possibly forming reservoir of infection, cannot be thoughtý, to a .. disseminate helminth infections.

Because of the greater significance of the-', former birds, '. ', were sought., additional sources of rooks, " jackdaws and starlings Mr. Peter Miller gave permission 'to trap rooks and jackdaws

Abbotshay Duck Farm, Ayot St. Laurence' and later- Smith,, '. at, b" . trap-, at Kline' and French Ltd., gave permission to erect a

Tallents Farm, Kimpton. In all some two hundred rooks and dissected from both farms. Starlings breed jackdaws were only ,.: in small numbers in Pembrokeshire and in the summer months of to the 1962 and 1963, more than five hundred carcases were sent ., writer by Mr. David Taylor of Blofield, Norwich.

Clapham (1940a), published the results of an experiment : she carried out at Winches Farm, St. Albans, in which the importance

of wild birds as disseminators of poultry helminths was stressed. As the facilities of Winches Farm were available, it was decided

to repeat Miss Clapham's work to determine any change in carrier

transmission which the more intensive methods of poultry farming

might have effected in the intervening years. The by the infection of selected 1, second approach was -wild, ,

birds with poultry nematodes to determine how much active opposed'; transmission of adult nematodes is possible as '. carrier , to passive transmission of eggs on external parts. The " birds

used were rooks, jackdaws, starlings and wood-pigeons, and,, the-_,

nematodes those shown in the survey to be the most common'-in: He ga Odin C. poultry, . Prsikiagallirarum, _a gal-21 and

tzSiglla a"

Initially, small suspensions of A- Pa1li and H. g;, 11in,, rnin

ova were supplied by Dr. Horton-Smith from Houghton. These were, ova,, embryonated at 28cC and fed to chickens. The, resultant '.°.. ý_ formed the basis of several 'subsequent infections... Mr. J. D. .,.. ; : 'Veterinary Blaxland at the Laboratory, New; Haw, Weybridge ', s., _ gave permission to use the facilities of, the Poultry Disease

Diagnosis Section and ,Dr. Kendal of the Parasitology Section_ also gave advice and some suspensions of Heternkf, Aýenrýdsýý, and C. Qhsig ntn ova. Ova suspensions were made by collecting adult worms from-, - the carcases sent to Weybridge for post-mortem purposes.. The, ' ; -, r- adult females of xlli were cut in half and the contents of`the , into bowl to being by hand pseudo-coelom squeezed a prior ground , --, with a pestle and mortar. The mixture through Ende-- was passed ,, . c ott sieves at meshes 100 and 150 and the ova were cultured, in

2% formalin in petri dishes 28CC. All 1, at specimens of ,gý1? any and oh dgnitri were collected with no attempt to separate males from females. These were ground and the mixture left todigest ..; in a solution of J% HCL and 1° pepsin to isolate the -After ova. . four hours in bottle 37S the was, a stoppered at , suspension passed through Endecott sieves at meshes 100,150 and 200. E attempt was made to differentiate the various species of C nil-

which may have been present since the resulting suspensions; were fed to the final host p= ,= and species such as n Infam which intermediate host require an would be automatically-": -: eliminated.

The cultured solutions were stirred daily to provide--fresh oxygen and the medium was changed every four days as a precautionýýý_ against concentration through evaporation. The method' of. collec- I 'were: tion included man infertile ova but sufficient embryonated ` obtained to develop strong infections in trial. tests on 12-day -`-,' . e day old, chicks. "The infective", doses administered ' consisted ova of two hundred to three hundred 'for 11l ýrtim and g 111

of which about one-third were infertile and four hundred -'ato ý. five hundred for nhcdg The higher dosage for i -i. o nig a 'n was used to conteract any, dilution through the presence of thei,

ova of n,-mdinfl, to or nnntig.

The rooks were maintained in an aviary at Winches Farm and.

fed on a diet of minced meat and grain. Birds were identified

by ringing as it was not possible to isolate individuals, for"

any length of time. The birds were trapped mainly in the ;winter, In late spring and early summer. summer and autumn, much,-'grain= was available in fields and the birds were not tempted-'into the-, traps. The experimental infections were divided into two periods corresponding to the availability of the birds. The first -,period the 1961 occupied summer and autumn of when birds were -obtained, '''_ from Abbotshay Duck Farm. Experiments through continued ,the winter, but in the following year the rearing of ducks was dis-', , ýý. continued and no new birds were available as rooks were-not, ',, . tempted into the traps in the absence of the ducks and. the, food. Nine birds from the trappings associated original were,,,, --- used through the summer of 1962. In March, 1963 the traps , at Tallents Farm were erected and fresh birds were available'

rooks and jackdaws from Tallents Farm were maintained -ý- on a Vitamin A deficient diet and successive infections, ',, were,, .., given to find out if resistance would be lowered by repeated doses. The ýdefic iency diet was that commonly used ' in, such work ,'., r and was recommended by Dr. D., ', Wakelin. _ The. aresz -ingredients i't110ý Wheat Oats '. 15, . , Wheat, middlings 25 Fish Meal 10 Unextracted Yeast 2.5 Lime stone Flour 2 Salt 0.5

with 1 million I. U. Vitamin D per ton. It was made by Sherrifs

of Welwyn Garden City.

In the winter of 1962, when it seemed doubtful whether more `;

rooks would be trapped, starlings wore caught in Pembrokeshire

and given infections of gý11 Four wood-pigeons were loaned ýý

to the writer and these were infected with 3H i and gi 1m in the same period. ý- RQatýlta '_'

Nnturn1 TnfAatntinns.

(i) Exafn inn hirds by . of yt1d Past-mortem. None of the wild birds examined in this survey e ontained ;

nematodes which are normally parasitic in domestic poultry, or. occurred Galliformes generally. Only Cn 1n nbn gntn, which ` l' in starlings, is also found in chickens and the turkey. On a

single occasion, a rook from a rookery at Ivy Towers, St. Florenea,

Tenby was found to possess a third stage larva which was' atypical ? (Zeder, 1800), the of Qrrnc, ýeým Pnsient?. d.Q . +im common ascarid Corvidae, it thought nematode of and on examination was _to'-be a larval A se ar 1cta spp.

(ii) Exnmirn. itinn of riir fromnxtnr_nni pnrts,

Clapham (1940b) has that -r- suggested soil attached _to"-the- ' feet and beak acts, as a medium for the mechanical, transport of_',', _ helminths evidence. from pasture but was unable ^to offer 'any Care was taken' to scrape all feet but "only -inthree birds, taken from Abbotshay_ Duck Farm were any ova recovered. These,: were a Heterakis spp. and were successfully embryonated. The eggs were then given p= ý. 5 to three two-week old cockerels but no adult worms were found when the birds were killed and dissected forty days. It is that the after possible species`,, -_.,. was d sp r (Schrank, 1790), which inhabits the caeca of, the -', goose and the-duck. Thus, there is unsatisfactory evidence, --, _---,,, that the rooks were capable of infecting pens placed on pre- viously uncontaminated ground at the farm and could easily ; transmit the to the ' nematodes any similar farm in vicinity.: -, (iii) The introductl: öd of nematodes to clean nastu_re, '

Clapham (1940a) reported the introduction of Cnpili. -aria = long ic o1 1 11, geterki s gr, 11 i narum, Tr iehoa_trnng, tAnIn i y1 is _a, Hym Asr1d1a ral i and norme; is orvi to young birds kept ' on ' <. plots which were formerly helminth free. This last species doubtful as the only cestode with this name occurs in rooks. in. ' the Southern United States and the writer has not been able to _ , ,, -_ trace the species as a synonym for any hymonolepid in the C or vidae of Europe. . A plot of ground was chosen at Winches Farm, which was sheltered and within ten yards of a copse. There had been: no; = __ poultry reared on this ground since at least the end of, the: -ýwar, so that it could be assumed to be free of holminth ova ý., from direct contact with chickens and any ova present would ,".. have-been" introduced by birds. In May,, 1962, arc was ý,. wild an , ' ,,T r ". 613 in the ten-day, :.. ' placed run and fourteen old chickens were ,-,, ;,. introduced together, broody. hen. The, faeces of the 'hen'',. ,, with'a were examined daily for six days and the bird proved-to be _"free, of helminth parasites. The chicks had been reared from eggs. in

a brooder.

At first none of the local fauna entered the run, and when-,

a poult was killed after one month, no parasites were recovered. - " (Table D 1). Later blackbirds, thrushes and house sparrows . ,, started to use the run. The second and third poults, also -`" " killed at monthly intervals, were negative. The fourth bird, , _. y. killed in September, was infested with three Heternkf s adult "`, gaits i nrum, The remaining birds were killed in successive.

months and the final eight at the beginning of December.,; ', 1n. -the, meantime, starlings were frequenting the run and rooks'. were

observed on adjacent trees. Ultimately large infestations ' with '; . fe aia were recovered with a maximum of 273 from one cock. -.--.:

A single bird was infested by six adult C_ nhýý nata. ;'';.

In 1963, the experiment was repeated on the same ground. - This was thought advisable since most small establishments have',

not the facilities to renew pasture yearly, more indication and ,= would be obtained the build infections. Again of up of young" . _', cocks were placed under the same broody hen. An ant-helmintic'

was administered to the hen but did not entirely remove>theý, ý

r infection. This was not considered important_ since

ova would be on the ground surface from the _-inevitably present ":_°' :r By, this time birds the, quite previous-year., wild -were using run 614 TABLE D. 1 ho increase ir. r_eriatode'_ infections, of poultry, placed on 'clean' pasture throuCh contact with wild birds. ('_:x-pre : eta as nurbcr of t: ormc per bird) . ,

ýcteral, a- l, ßc ridia Capillaria, Canil1aria 1 96, I' rd ,-a1'inarura Zalli obaiCnita caudinflata

June 1 - - - -

July 2

August 3 - - - -

Sept. 3 - - ,-.. Oct. 5 - - - _.

Fov. 6 31 - _ -,

Dec. 7-14 97 - 6 (one bird only) x: fimures - Two girds uninfectee, The given represent the averaGo ,. nuMber of worix preoent in the six infected birds.

1963

June 1 11 3 - e_7 July 2 15 28 '11=- - , August 3 23 3

Sept. 4 53 4 30

Oct. 5 69 9 45 22ý

6 Nov. - 2 31 x - Dec. 7-12 71 5 42 61 freely and only rooks: were, rarely'seen to alight within the ° boundary. Sparrows' flew, in, from a wire-netting poultry, -run at, the far end of the farm.

The first bird of the second series was infected with, ü_. it birds found to _ra and subsequent were contain Annnridi, i, ma- C. ohs nl and C. r anjinf] . a. No cestodes -or, tre todes were recovered. Since nematodes could be introduced--by

mechanical transmission, the lack of cestodes could be-, attribute'd',

to either the lack of a source of infection for the wild. birds

or to the absence of a suitable intermediate host. The . infec tion with C_ ermdinfl ti was slight and had died out by Christmas. '

The infection is most likely to have been introduced via infected:, have in the faeces the worms, parts of which would passed out :ýof- -l wild birds. Experimental work was discontinued in January, ", 1964, `. and it was not possible to find out whether the earthworms °in ', _ the run became infected. In this second period a definite- ' _ý _. seasonal incidence was found which reached a peak in Sept enber.,. ",- and October. The bird killed in October was the only one to', be infested by all four species of nematode.

B).

The series of forty-two experiments is listed consecutively, '.. in Tables D2 3. and r: (i) gen s mnin ni and nn a normal dint (Julys 19b1-Novemb©rý 1962).

Altogether there were thirteen attempted infections, '-five with Heterakij (Nos. 1,4,7,9,12), five with 69, 8,114) 'and 111 ia ýt (Nos. - three with C an nr obs ý 3' 1O1 16) 1U mair: Table . 2: Experincntal infections of Sire ".ainea cri normal `U die xz. (Jul 1961 » January, ,, -1963) - Hetoraki:. E Cýt7iýf. t , ccaridiý ar a. r-all! :ýn: ur3 r-r31i aýýýi r°t ±

1 Rook (1)

2 rook (1)

3 nook (1)

1F z ookk (2 )

5 : 1ook (a )

5 nook (3)

7 : oolý (3 )

8 :took (4)

9 rook (4)

10 Rook (2)

11 Pigeon (1)

12 Rook (5)

13 Pigeon (1)

14 Rook (5)

15 PiCeon (2)

16 Rook (3)

17 Pigo (2)

18-21 Starling (1-4) 6 t7 In the first half, dozen experiments, -six birds were : infected, killed after eight -and for, worms.. (Ackert weeks-, examined adult ý 1937, showed that A. Pallf takes fifty days to attain-,, maturity) There followed a dearth of birds and the same rooks were re-, -', infected every four weeks from March, 1962 to November, ` 1962. (Expts. 7,8,9,10,12,14,16). After experiment 16, six , birds killed were and examined for adult worms, the other,, three '_ý- birds having died in the meantime. The control birds for infection ý:. used each were chickens, -,.! -° maintained in parasite-free conditions at the Animal House of''.,

the London Schbol of Hygiene and Tropical Ihdicine. The'-, via- bility of each suspension was proven by the successful-establish-l' ment of adults in these birds. Not it unexpectedly, proved impossible to infect rooks ,,with '',- He ernkInt. The normal habitat in the chicken is the caeca and only in super-infestations are the nematodes regularly found; in the adjoining regions of the small intestine. It was thought z_ that the infection of an un-natural host, which has only- very. _

small caeca about an 1" long, might induce the development -. of '-` , --'. : the nematodes in abnormal regions of the small intestine but this did not prove to be so. Fresh faecal samples from the-rooks"

collected twenty-four to forty-eight hours after infection, con- `, tained the unhatched ova to produced which, when given chickens,,,, __""' infection. an Thus the rook cannot act as host to Heterikisbut, I., can passively transport the ova in the intestine. ý = In contrast examination of faeces of rooks, infected,, with_,

Aqcnr jda gam that 75 the hatched and, first _, -.. showed of' ova stage larvae were present. One bird was ' isolated and, killed `, -` twelve after, days, by mich time, in the chicken' the duodonal °

mucosa would have been penetrated by the larvae. There were no,,

larvae present and no signs of penetration. It is concluded

that while rooks under special conditions might act as host to

adult AQc0 fdf they are less likely to spread infections of

1ýscýrr than of HP er k1 because fewer ova pass intact through

the anus and it is known that chickens rarely become infested: by ingesting larvae to This is in Table D as opposed eggs. seen _'1" where A- gal-Ti was the least prevalent intestinal nematode.

The third series of infections with (nomii grin ohygn tý

was successful and adults were recovered in small numbers. -

(Experiment 16). As adults had not been established in, -the might previous experiments, the possibility arose that the diet

have modified intestinal conditions over a period of months. All. " , birds were infected with Q- rnnanta (Dujardin, 1843) and `several.. Po iej (Zeder, 1800). (Established by with rocne - um enn in. am 'after,, -, faecal analysis). The ansicnuda . 1m infection disappeared three but the G, infection was retained about months, __reSe _ta -throughout the period, so that the diet might well have been-

more favourable for infestation with C pfll ria spp. Rooks can be r. Ova coven- considered to be active carriers of - ohsi sta. trated from the rook faeces were infective when fed to'chicks., (1960) the'- Stoiean fed ova of ""0- - rvorum"" obtained=from intestines of rooks to chicks and was able to effect an infection. - (Rudolphi, 1819) to'; be Most authorities considered CC. ornattiýi _. description in the, a-, omen ridum and there is no relevant ; -liter 6 tore. The' species recovered by Stoican may have been c_ (Wassilkowa & Gouchanskaja, 1930), 0 (if : cgwicfli_ -.. rem .: this is ignit Experi-.. species a synonym of anti) or -ob . mental evidence in this thesis and that of Wakelin (1963) host Stoican', s suggests that n . ia requires an intermediate and species is most likely to have been 0bs1cwta. (ii). 1 (Sept. 1962-Dec. 1962). ° Pi-en cmn. gined on a norm diA . Miller (1937) was unable to infect pigeons with Hatera1d = or A-qgaridia. Four wood-pigeons which had previously been -used :, for bird loaned to the writer. While =ý malaria experiments were , .:, their importance as disseminators of poultry nematodes is-minimal-l', since wild pigeons rarely move in the vicinity of dwelling they tolerate: =ý places, it was of interest to see whether would "_- H idi they frequently infections of nrnk and As -a as are by Q. C. langico i (Mehlis, lß31) parasitised obsig a .-a and =a parasite of game birds. There were two attempted infections,. öý _, , (Expts. 11 and 15) with Hrt rnkla and two with Ase ridi '(Expts.. ,. "_ "ova 13 and 17). Neither species established itself and the : -.: passed in the faeces were still viable. '-.- (iii) Starling mnintatned on o nn1 d At. Q)ec. 1962-Jan. 1963).

Six kept in trap in Pembrokeshire: The starlings were a ... small size of the caeca would preclude development of }{et rakig -ý_ and it seemed unlikely that so large a worm as be tolerated. Of all species of birds, the starling isthe, most'-,. ý.. suspect in the dissemination of disease and super- infestation,, ý. with A- gall, was attempted by administering four. ýweekly infec- tions. Predictably,,. developed and the - ova, which were no . adults , . 6 20, passed-in the faeces withintwenty-four hours were taken, to. --. London infective to chicks. fe and proved when, given _oob "a has been from the infection, -: recorded starling and experimental , -, was superfluous. (iv) (March, 1963-November, 1963).

There are many reports of an increase in the number, and, size of nematodes infesting fowl maintained on deficiency a ,., .` diet. As it had been possible to infest rooks with C. 'ohLignnt4

and demonstrate a partial toleration for AT, when rooks

and jackdaws became available from Tallents Farm, it wash decided

to determine whether the ability of the nematodes to establish ',

themselves in these birds would be increased in combination with deficiency diet. The infections is in Table-, ', a sequence of seen -, D 3. It was not possible to separate the two species 'of. birds of` in the aviary so care was taken to infect only one species -ý ; bird with a particular species of nematode at any one time., Thus took _ experiments 22 and 23,25 and 26,27 and 28 etc. , place simultaneously with a four week interval between each- ;,

successive dosage. As before, control chickens were roared in, the Animal House at the London School of Hygiene and Tropical Medicine. birds infected time butt Initially, six wore at one - ,,, showed--that towards the end of the period, odd birds died and lesions :

death was due to the deficiency.

There experiments, three for each host with were sixteen ,: G.ý A g:,-, rte and HPtarakt s, and two for each host with ' ohsig A. and, für larger '% This ' last species established itself in both birds,, Table 624 fl.: Experimen tal infections of -Dirc1o maintained on _vittamin A, (;: 63) deficient diets. arch, 1 963 Cctober, 1 :-.

(N. ß. The infection of duck: with, C.. rececta is , incl uded in the.. ': sequ ence- for, conven ience. he birdo were, of `cours e, " maintained on ) a no rmal diet. -;

! Heterakis A:, caridia Capillaria Capillaria Expt* mal 3 inarun rx7.1i obsiEnatn rececta

22 Jackdaw (1)

23 ? ooh (1)

24 Duck., (1)

25 Jackdaw (1)

26 Rook (1)

27 Jackdaw (1)

28 Rook (1)

29 Jac}: d, w (ý )

30 Zook (2) 5

31 Duck 2)

32 Jacltdaw (')

33Z oos- (2)

34 '?ook_ (2)

35 Ducý. (3)

36 Jackdaw (3) (3) 37 -took. ) 38 J)uckc(f,

39 Jackdaw (3)

40 Rook (3)

41 Jacl. dav (2)

42 Duck (5) 622, on numbers in the rook than when the bird was; 'r©ared- a normal dißt. Heeterakis was: 1n host. '..:' The -not established either, Ascar idin. ova hatched and immature adults were established '.in The intestinal 'the, later infections. lesions resulting from _ :,-.: deficiency lowered the host's resistance and made it possible, for the larvae to develop in the mucosa. The birds died, or were killed before the adult stage was reached but corvid' birds can evidently be infected with A_ g l1i. In the field such birds are most unlikely to be deficient in a factor,, Such,,, '. as Vitamin A and it must be concluded that wild birds do not _. act as hosts for the more common poultry nematodes with. the exception of ph4fgrL1th. slimmnr

1) birds to be transmitting, the _' . Wild are shown capable of ova of poultry nematodes. The most probable method is by ingesting the ova either in faeces or in infected earthworms, which pass,,, ', unharmed through the intestine. This is more true for Hetnrnk g- than Asen idia which can hatch in the intestine of certain, corvid birds. transmit 'Passive` 2). Wild birds C-npi? rin oh4ig n t^ both by transmission of ova and by acting as hosts to small numbers of..: adults. The importance birds infections is, ' of wild as carriers of - ,- limited to free range poultry and those intensive systems to ', which they have access. The poultry in such units will -usually have contact the domestic ' . only with small passerines such as sparrow, which is indigenous to a particular farm or dwelling

place. It is possible that sparrows might, carry ova from the

faeces of larger birds into units but evidence could only be ,"y provided with difficulty and is not offered in the present", work.

r` APPS The life-cycle Cnp i11nria (Dujardin, of ran . _tn . 1843), parasitic in the small intestine of the Rood (Corv«s frtigf L. ).

The rooks trapped at Abbotshay Duck Farm were 100% 'infested',

with C. Z eta. Their morphology is described in Part " 1j

Section C. The principle interest lies in the proximity of ,the'`ý

rookery and the duck farm and the fact that certain authorities,, notably Madsen (1945) consider scse a to be a synonym, of 'an t1

(Schrank, 1790), which inhabits the small intestine

duck and various domestic and gallinaceous birds. The question änd '. arose whether the species could be proved to be synonymous " ' ,".

whether the rooks were infected by the ducks or vice vaza. -_ cross-infections could be obtained then the small morphological:. ',."

differences such as egg shell pattern and shape might prove "to

be due to change of host. In an instance such as this,, where

both hosts ate the same diet, Le. the duck food, then -physio- '1

logical adaptation might be partly negated and cross-infections more easily obtained. It was decided to attempt cross-infection to determine the necessity or not of an intermediate host. and ., However, the examination of many duck intestines which-, were -s-ý-. disposal, the `;. -placed at the writer's showed complete' absence_of: C restricted. to 'the ,"ý_ any piljnr n spp" , and experiments were ,,,, 62' attempted infection of ducks with ' ova obtained from the ' , , rooks. Nat Mf?thods _r ;r1.1 and The nematodes were collected from the rook intestine and the isolated described for in, _, ova and embryonated as oeaignntnn ,. the introduction to Part IV. By the time the for `.: preparations. . the experiments were complete, the duck farm had closed and no further rooks were available to act as control birds until March, 1963. These latter birds were less frequently Infested- with rp and the isolation of several, followed by faecal examination at daily intervals, resulted in six 'clean', birds , -. , being used as controls in the experiments.

The ducklings were reared in parasite free conditions and. were kept in the Animal House at the London School of Hygiene at' and Tropical Medicine. The control birds were also kept', - the Animal House. Two methods of infection were used; direct,.;

indirect via intermediate host. The ,Q. and an annelid earthworms the farm identified A11o "eä1 ig at were as ohophor, ri.. . Elsa fn tEir1 by Mr. R. W. Sims, Section, . and nin . Annelida British Museum.

8u]L

The sequence of five experimental infections are--shown in Table D 3. (Experiments number, 24,31,35,38,42) ._. (i). Direct infection pis

In experiments 21+ and 31, eighty to one hundred ýandä.fifty : embryonated ova were given to each of six three week: `old duck-", lings.; ' given. 'to ones control`ý, -'A single, infection of ova was. _ bird. -, After, four weeks the control bird was killed and adult The rt were- recovered. , faeces -of the' ducklings, 'were

analysed daily for ten days and were consistently negative.

After experiment 31, the ducklings were killed and examined <-ý for adults but none ware present.

(ii). Indirect if ion thron hwjarm, o? . . vjr !p_ _ies ör en If r to and imn tis are synonyms, then while the `t

form was shown in the previous two experiments to have a ýdirectý. ý

life cycle, it is possible that the nnatIR form may require

intermediate host. Three of the more common genera of earth-

worms were collected and used in consecutive experiments, :: _ x Ai in 35,1A 1 in änd_ enia experiment 01 ,"bophnrr experiment 38, .ýr L imbri ig in experiment 42. The worms infected `sixty _ were withý. ,:. to eighty ova which were introduced into the buccal cavity 'byý, ý ewers-, a very fine pipette. After twenty days, six earthworms ý'. given to each of six ducklings and a further six to the control.:

bird. The control bird was killed four weeks later in, all-, '-- experiments in 35 38 the duckling faeces examined. `,, and and were - At the conclusion of experiment 42, the ducklings were killed-, ',,, and dissected.

None of the ducklings infected the bird were and control, . :°ý. was infected only via Aflo1nhn phor , the more common of the two species at the duck farm. =

C;one I nie s

C iiinrir rP, et, (Dujardin, 1843) has a direct : life= ý- cycle. Unless intermediate host is t -, . an unusual required, - -',

62

The writer wishes to thank Professor J. J. C. Buckley.: and Dr. C. Horton-Smith for their respective supervision of ý:, the taxonomy and ecology of the wild bird helminths, T and ý. the survey and experimental work with poultry helminths.

The greater part of the work was carried out under,, the tenure of a grant from the British Egg Marketing Board and much encouragement was given by Dr. N. R. Knowles, Director; -, of the Scientific and Technical Division. A grant for: equipment used at Winches Farm was given by the Governors, ''_, '-, of the London School of Hygiene and Tropical Medicine. ý, Technical advice and assistance was provided by,.',. ý_

Mr. F. R. N. Pester, W. B. L. Cheetham and It. E. Blackie'., of the London School of Hygiene and Tropical Medicine and by-

Its Blaxland and his colleagues at the Central Veterinary,,; " Laboratories, Weybridge. Help in the identification' of- species was provided by 2r. S. Prudhoe of the Natural History -,

Miseum and by Dr. D. Wakelin and the plates were prepared the Photographic Department of the L. S. H. T. M. Finally, to Miss ý; ' the writer is grateful A. Witheeombe, -. and Mrs. R. Eversden for typing the text and to Mrs. S"; Buck: for preparing the tables. REFERENCES ". "

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