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The taxonomy and biology of splendidofilariine nematodes of the tetraonidae of British Columbia Gibson, George Gordon 1965

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The U n i v e r s i t y  of B r i t i s h  Columbia  FACULTY OF GRADUATE STUDIES  PROGRAMME OF THE FINAL ORAL  EXAMINATION  FOR THE DEGREE OF DOCTOR OF PHILOSOPHY of GEORGE GORDON GIBSON B.A.j  University  o f Toronto,  1955  M.A., U n i v e r s i t y o f T o r o n t o ,  19 57  THURSDAY, AUGUST 5, 1965, AT 10:00 A.M, IN ROOM 3332, BIOLOGICAL SCIENCES BUILDING  COMMITTEE IN CHARGE Cha i rman: I.McT. C owan J . R. Adams J . F. B e n d e l l J. B i e l y External  K. Graham D. C. G. MacKay M. D. F. Udvardy  Examiner: M.B. Chitwood  P r i n c i p a l Research P a r a s i t o l o g i s t , B e l t s v i l l e P a r a s i t o l o g i c a l Laboratory, U.S. Department o f A g r i c u l t u r e  THE TAXONOMY AND BIOLOGY OF SPLENDIDOFILARIINE NEMATODES OF THE TETRAONIDAE OF BRITISH COLUMBIA ABSTRACT T h i s study was undertaken t o determine the i d e n t i t y of the c a u s a t i v e agents of the f i l a r i a s e s r e p o r t e d from northwestern t e t r a o n i d s (Fowle, 1946; Babero, 1953), the i n c i d e n c e of f i l a r i a l i n f e c t i o n s i n the T e t r a o n i d a e of B r i t i s h Columbia, and the means by which these b i r d s acquire t h e i r i n f e c t i o n s . More than 400 w i l d g a l l i n a c e o u s b i r d s of 10 s p e c i e s and over 200 a d u l t n o n - g a l l i n a c e o u s b i r d s from 27 f a m i l i e s were examined. A d u l t s of the f o l l o w i n g 4 s p e c i e s of f i l a r i a e (Onchocercidae: S p l e n d i d o f i l a r i i n a e ) were c o l l e c t e d from t h e T e t r a o n i d a e of B.C, and/or Ala-ska and a r e d e s c r i b e d h e r e i n : 1) S k r j a b i n o c t a f l e x i v a q i n a l i s (Jones, 1961) n. comb.; 2) S p l e n d i d o f i l a r i a p e c t o r a l i s new s p e c i e s ; 3) S p l e n d i d o f i l a r i a " s p e c i e s A"; and 4) Splendidofilaria p a p i l l o c e r c a (Lubimov, 1946) Anderson and Chabaud, 1959. D e s c r i p t i o n s a r e p r o v i d e d a l s o f o r the m i c r o f i l a r i a e of S f l e x i v a q i n a l i s . j>. p e c t o r a l i s and _S. p a p i l l o c e r c a , and f o r M i c r o f i l a r i a l a g o p o d i s (Haaland, 1928) Brinkmann, 1950, and M i c r o f i l a r i a " s p e c i e s B" from tetraonid hosts. Recent systems of f i l a r i o i d c l a s s i f i c a t i o n a r e e v a l u a t e d and some m o d i f i c a t i o n s proposed, i n c l u d i n g r e v i v a l of the genus JSkr. j a b i n o c t a Chertkova, 1946 t o comp r i s e _S_. p e t r o w i Chertkova, 1946, S_, f l e x i v a g i n a l i s , j5. chitwoodae (Anderson, 1961) n, comb., S, s t r i a t o s p i c u l a ( H i b l e r , 1964) n, comb,, and p r o v i s i o n a l l y S, l i e n a l i s ( O r l o f f , 1947) n, comb. On Vancouver I s l a n d , M i c r o f i l a r i a sp. B i s hyperenz o o t i c i n b l u e grouse, w i t h a s i g n i f i c a n t l y h i g h e r prev a l e n c e i n a d u l t males than i n a d u l t hens or i n y e a r l i n g s (or c h i c k s ) . M|_, sp, B i s e n z o o t i c i n Vancouver I s l a n d r u f f e d grouse, J3. f l e x i v a g i n a l i s i s h y p e r e n z o o t i c i n Vancouver I s l a n d b l u e grouse; i t s p r e v a l e n c e i n a d u l t males, and i n a d u l t females i s s i m i l a r and i s s i g n i f i c a n t l y h i g h e r than i n y e a r l i n g s . _S. f l e x i v a g i n a l i s i s sporadic i n B r i t i s h Columbia r u f f e d grouse. Splendidof i l a r i a p e c t o r a l i s i s sporadic to enzootic i n tetraonids from c e n t r a l B. C. and A l a s k a . Mf. l a g o p o d i s i s e n z o o t i c i n w i l l o w ptarmigan of n o r t h e r n B. C, M i c r f i l a r i a e c l o s e l y resembling Mf. f l e x i v a g i n a l i s o c c a s i o n a l l y  p a r a s i t i z e n o n - g a l l i n a c e o u s b i r d s of Vancouver I s l a n d , but Mf. sp.B and J3_. p e c t o r a l i s seem to be r e s t r i c t e d to tetraonid hosts. The l a r v a e of ^S_, f l e x i v a g i n a l i s develop i n the t h o r a c i c muscles of the c e r a t o p o g o n i d , C u l i c o i d e s u n i c o l o r (Coq.) group. Those of Mf. sp. B develop i n abdominal f a t b o d i e s of the s i m u l i i d s , Simulium aureum F r i e s and Cnephia minus (D. and S.); the former being c o n s i d e r a b l y more e f f i c i e n t as an i n t e r m e d i a t e h o s t . Development of Mf. sp.B f a i l s at temperatures which .do not range above 62°F. L a r v a l development i s d e s c r i b e d and on the b a s i s of l a r v a l c h a r a c t e r s some f e a t u r e s of the a d u l t s of M f s p „ B are p r e d i c t e d . Knowledge a c q u i r e d on the b i o l o g y of the t e t r a o n i d h o s t s , the i n c i d e n c e of the f i l a r i a s e s , and the seasonal abundance of the v e c t o r s and the e f f e c t s of v a r i o u s f a c t o r s on t h e i r f e e d i n g a c t i v i t i e s i s i n t e g r a t e d i n a d i s c u s s i o n of the e p i z o o t i o l o g y of Skr j a b i n o c t a f l e x i v a g i n a l i s and M i c r o f i l a r i a sp.B,  GRADUATE STUDIES Field  of Study:  Parasitology  B i o l o g i c a l Methods and Procedures P r i n c i p l e s of W i l d l i f e B i o l o g y and Conservation B i o l o g y and P o p u l a t i o n Dynamics of G a l l i f o r m B i r d s Advanced P a r a s i t o l o g y Marine F i e l d Course Other  Zoology S t a f f 'I, McT,  Cowan  J,  F, B e n d e l l J , R. Adams P. A. Dehnel  Studies:  P r i n c i p l e s of P l a n t Ecology P r i n c i p l e s of C o m p a r i t i v e Psychology P o u l t r y Diseases and Hygiene  V, D. J.  J. Krajina  C. G. MacKay E. L a n c a s t e r  PUBLICATIONS bson, G. G. 1964.  Taxonomic and  o b s e r v a t i o n s on S t r e p t o c a r a  biological  californica  ( G e d o e l s t , 1919) G e d o e l s t and L i e g e o i s , 1922 and the genus S t r e p t o c a r a (Nematoda Acuariidae).  Can. J . Z o o l . , 42: 773-783  THE TAXONOMY AND BIOLOGY OF SPLENDIDOFILARIINE NEMATODES OF THE TETRAONIDAE OF B R I T I S H COLUMBIA by GEORGE GORDON GIBSON B.A., U n i v e r s i t y M.A., U n i v e r s i t y  o f T o r o n t o , 1955 o f T o r o n t o , 1957  A THESIS SUBMITTED I N PARTIAL FULFILMENT OF THE REQUIREMENTS FOR,THE DEGREE OF DOCTOR OF PHILOSOPHY in  the Department of Zoology  We a c c e p t t h i s required  thesis  as c o n f o r m i n g  t o the  standard  THE UNIVERSITY OF B R I T I S H COLUMBIA A u g u s t , 1965  In p r e s e n t i n g the  this  thesis  Columbia,  I agree that  the Library  a v a i l a b l e f o r r e f e r e n c e and s t u d y . mission  f o r extensive  p u r p o s e s may  of this  w i t h o u t my w r i t t e n  Department o f  It i s understood  thesis  for financial  permission.  Zoo l o g y  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, C a n a d a  August,  1965  thesis  by t h e Head o f my  Columbia  per-  for scholarly  D e p a r t m e n t o r by  that  gain  of  s h a l l make i t f r e e l y  I f u r t h e r agree that  copying o f t h i s  be g r a n t e d  representatives,,  cation  Date  fulfilment of  r e q u i r e m e n t s f o r an advanced d e g r e e a t t h e U n i v e r s i t y  British  his  in partial  copying or  shall  publi-  n o t be a l l o w e d  Chairman:  P r o f e s s o r James R. Adams  ABSTRACT  This study i d e n t i t y of the reported  agents of the  from northwestern t e t r a o n i d s  Tetraonidae  which these  incidence  b i r d s acquire  s p e c i e s and  over 200  and  of f i l a r i a l  the means by  w i l d g a l l i n a c e o u s b i r d s of  Adults  of the  (Onchocercidae: Tetraonidae  are d e s c r i b e d h e r e i n : 1961)  10  b i r d s from following 4  Splendidofilariinae) of fi, ".C. and/or  l) Skriabinocta  flexivaqinalis  (Jones,  p e c t o r a l i s new  s p e c i e s ; 3) S p l e n d i d o f i l a r i a  and 4)  1946;  infections in  adult non-gallinaceous  were c o l l e c t e d from the Alaska  (Fowle,  their infections.  f a m i l i e s were examined.  s p e c i e s of f i l a r i a e  the  filariases  of B r i t i s h Columbia, and  More than 400  27  undertaken to determine  causative  Babero, 1953), the the  was  n. comb.j-2) S p l e n d i d o f i l a r i a "species  A";  Splendidof i l a r i a p a p i l l o c e r c a (Lubimov, 1946;)  Anderson and  Chabaud, 1959.  a l s o f o r the m i c r o f i l a r i a e p e c t o r a l i s and laqopodis "species  D e s c r i p t i o n s are of S.  flexivaqinalis.  S. p a p i l l o c e r c a . and  and  Microfilaria  from t e t r a o n i d hosts. Recent systems of f i l a r i o i d  evaluated  S.  for Microfilaria  (Haaland, 1928),Brinkmann, 1950, B"  provided  and  classification  some m o d i f i c a t i o n s proposed, i n c l u d i n g  r e v i v a l of the genus S k r i a b i n o c t a Chertkova, 1946  to  are  comprise S.  S.  petrowi  chitwoodae  Chertkova,  (Anderson,  1961)  (Hibler,  1964)  n.  comb., and  (Orloff,  1947)  n.  comb.  On  Vancouver  hyperenzootic prevalence yearlings Island  i n blue  n.  Mf.  grouse.  Columbia  S.  ruffed  comb,, S.  with  sp.  grouse.  ptarmigan  resembling  Mf.  S.  The  seem t o be  l a r v a e o f S.  thoracic  muscles  o f the  unicolor  (Coq.) g r o u p . fat bodies  Fries  C n e p h i a minus  and  considerably Development  o f Mf.  range  and  on  the  a d u l t s of Mf.  the  (D.  more e f f i c i e n t  not  C.  is.enzootic  Microfilariae  sp.  above 62°F. b a s i s of  sp.  restricted  closely  and as  S.); an  develop  B develop  Simulium  the  former  intermediate  at temperatures  L a r v a l development characters  B are p r e d i c t e d .  sp.  in  B  the  Culicoides  sp.  simuliids,  Mf.  t o t e t r a o n i d hosts.  flexivaginalis  B fails  larval  in  occasionally parasitize  ceratopogonid,  of t h e  in  i n t e t r a o n i d s from  Those o f Mf.  abdominal  higher  i n Vancouver  b i r d s o f V a n c o u v e r I s l a n d , but  pectoralis  B is  i s sporadic  lagopodis  B.  flexivaginalis  non-gallinaceous and  Mf.  of n o r t h e r n  sp.  Splendidofilaria  C.  Alaska.  lienalis  a significantly  flexivaginalis  central willow  S.  B i s enzootic  i s sporadic to enzootic and  striatospicula  i n a d u l t hens o r i n  pectoralis B.  flexivaginalis.  Island, M i c r o f i l a r i a  grouse,  or c h i c k s .  S.  provisionally  i n a d u l t males than  ruffed  British  1946,  in  aureum being host. which  do  i s described  some f e a t u r e s  of  iv Knowledge acquired on the b i o l o g y of the t e t r a o n i d hosts, the i n c i d e n c e of the f i l a r i a s e s ,  and the seasonal  abundance of the v e c t o r s and the e f f e c t s of v a r i o u s f a c t o r s on t h e i r feeding  activities,  i s integrated i n a discussion  of the e p i z o o t i o l o g y of S k r i abinoct a f l e x i v a q i n a l i s and Microfilaria  Examiners:  sp. B.  V TABLE  OF  CONTENTS  Page GENERAL PART  INTRODUCTION.  ONE.  1  MORPHOLOGY AND  TAXONOMY  INTRODUCTION MATERIALS RESULTS I.  6  AND  AND  METHODS  2.  B.  SECTION  F i l a r i o i d s 1.  2.  3.  10  of  . . . . . .  Cryptic  Skriabinocta  Description  b.  Summary  c.  Discussion  Sites  14  of  14 14  occurrence  17 17  Splendidof i l a r i a a.  Description  b.  Summary  c .  Comments of  14  flexivaginalis  a.  F i l a r i o i d s 1.  .  DISCUSSION  DESCRIPTIVE A.  .  of  sp.  A  •  27 .  occurrence'  .  27 28 .28  Subcutaneous  Splendidof i l a r i a a.  Description  b.  Summary  c .  Discus sion  of  Sites  pectoralis  n-.  29 .sp  29 29  occurrence  32 33  Splendidof i l a r i a  papillocerca  34  a.  Description  .  34  b.  Summary  c.  Discussion.  Comments  on  of  S.  .  .  occurrence  34  . papillocerca  35 and  S.  pectoralis.  35  vi TABLE OF CONTENTS  (continued) Page  C.  M i c r o f i l a r i a e of Unknown A d u l t s . 1. M i c r o f i l a r i a  laoopodis.  a. Comparison with  . . . . . . . . .  44  .  44  other d e s c r i p t i o n s  44  b. Comparison with Mf_, p a p i l l o c e r c a 2. M i c r o f i l a r i a  sp. B. .  45 . . .  48  .  48  a. D e s c r i p t i o n . b. Summary of occurrence.  . .  c. D i s c u s s i o n I I . TAXONOMIC SECTION. . . . . . . . . . A.  49 . . .  49  . . . .  53  .  53  C h a n d l e r e l l a . S p l e n d i d o f i l a r i a . and Related Genera. 1. H i s t o r i c a l  summary and s i g n i f i c a n c e of  c u t i c u l a r bosses  B.  2. The genus S k r i a b i n o c t a Chertkova, 1946. . . . .  60  3. The genus C h a n d l e r e l l a .  64  4. The genus S p l e n d i d o f i l a r i a .  73  Comments on Recent Systems of C l a s s i f i c a t i o n of the  PART TWO.  53  Filarioidea  . . . . . . .  . . . .  78  BIOLOGY  INTRODUCTION. .  86  MATERIALS  90  1. Study Areas. 2. D i s t r i b u t i o n , Habits and Habitat Preference of G a l l i n a c e o u s B i r d s . . . . . . . . . . . . . . . . 3. I n f e c t i o n s i n Grouse Used f o r Exposure  90 93 100  vii TABLE OF CONTENTS  (continued) Page  METHODS  ; ,  102  1. C o l l e c t i o n of Data on N a t u r a l I n f e c t i o n s . 2. Determination  . 102  of M i c r o f i l a r i a ! P e r i o d i c i t y  102  3. Maintenance of Captive Grouse  103  4. Exposure of I n f e c t e d Grouse and C o l l e c t i o n of B i t i n g F l i e s  104  5. Maintenance of Engorged F l i e s  106  6. Developmental Stages  107  7. Experimental  . . . . . . . . . .  Transmission.  .  108  8.. C o l l e c t i o n of Data on N a t u r a l Transmission  110  RESULTS AND DISCUSSION. . . . . . . .  I l l  I. DISTRIBUTION AND PREVALENCE OF NATURAL FILARIAL INFECTIONS A.  I l l  Incidence of F i l a r i a s e s i n G a l l i n a c e o u s B i r d s .  . .  1. Occurrence .on Vancouver I s l a n d  Ill 112  a. Prevalence  of S k r i a b i n o c t a f l e x i v a g i n a l i s .  . 112  b. Prevalence  of M i c r o f i l a r i a  . 113  sp. B . . . . .  c. Comparative prevalence of these 2 species on Vancouver I s l a n d 2. Occurrence of i n f e c t i o n s i n c e n t r a l and northern B r i t i s h Columbia and Alaska a. Incidence of S k r i a b i n o c t a f l e x i v a g i n a l i s b. Incidence of M i c r o f i l a r i a  sp. B  c. Incidence of S p l e n d i d o f i l a r i a p e c t o r a l i s d. Incidence of M i c r o f i l a r i a  115 . . 116 116 . . 116  lagopodis  (?-S. p a p i l l o c e r c a ) \ \ i n Non-Gallinaceous  113  . 117  B.  Incidence  Birds  C.  Summary and T e n t a t i v e Proposals  120 122  v i i i  TABLE  OF  CONTENTS  (continued) Page  II.  OTHER  INFORMATION DERIVED  INFECTIONS  FROM  of  On  Vancouver  2.  In  the  Indications  of  C.  Longevity  D.  M i c r o f i l a r i a l  E.  Summary  of  and  Region.  Prepatent  A. B. C.  OF  .  .  .  .  .  .  Period  Infections.  .  .  .  .  .  .  .  .  .  .  .  Periodicity  Conclusions  FILARIAL  Relative  of  Simuliidae. D.  Influence  E.  Comments  .  of on  2.  Development  LARVAE  ORNITHOPHILIC A. B.  sp.  B  136 140  in  .  143 on  Development.  . . . .  Development  150 153 153  as  an  i n d i c a t o r of  systematic 154  DIPTERANS  . 1 5 8  Haematophagous D i p t e r a n s Study Areas Grouse-Biting Insects as and  128  136  position V.  .  135  Microfilaria  Larval  General  127  133  . -  Temperature  1.  .  to  Fourth-Stage Larvae of S p l e n d i d o f i l a r i a p e c t o r a l i s n. sp Development of S k r i a b i n o c t a f l e x i v a q i n a l i s in Culicoides Development  124  129  "  TRANSMISSION.  DEVELOPMENT  123 123  Transmission  IV.  Period  Island'  Okanagan  B.  123  Transmission  1.  ARTIFICIAL  FILARIAL  -  A. . I n d i c a t i o n  III.  NATURAL  Vectors  1.  Mallophaga  2.  Tabanidae  C o l l e c t e d from Intermediate  the  Hosts  '  158 160 160  . . . . .  160  ix  TABLE  OF  CONTENTS  (continued) Page  3.  Culicidae  •  4.  Hippoboscidae.  5.  Simuliidae  as  hosts  for  6.  Culicoides  as  hosts  for  flexivaginalis. 7. C.  161  Mf.  sp.  B.  .  .  .  .  .  161  .  162  .  165  Skriabinocta  .  Summary.  Influence  167  of  Exposure  Duration  on  Collection  Efficiency  D.  E.  167  1.  Collection  of  simuliids  2.  Collection  of  Culicoides  Seasonal  Fluctuations  1.  Trout  Creek.  2.  Nanaimo  Feeding  of  Vector  Lakes  Activity of  169  .  170  172 of  Vectors  Relation  2.  and h a b i t a t . . . . . . R e l a t i o n of a c t i v i t y to  a c t i v i t y  to  174 structural  factors  . . . . . . . . environmental  .  .  174 178  EPIZOOTIOLOGY  183  CONCLUSIONS  LITERATURE  CITED  TO  END  LITERATURE  CITED  IN  PART  APPENDIX.  .  .  170  1.  SUMMARY AND  Populations  .  factors. VI.  168  194 OF  PART  TWO  ONE  199 206 211  X LIST  OF  TABLE I.  II.  III.  IV.  V.  VI.  VII.  TITLE  IX.  X.  XI.  21  Morphometric; d e s c r i p t i o n s of Females of S k r i a b i n o c t a f l e x i v a q i n a l i s and s i m i l a r species  23  Morphometric descriptions of S k r i a b i n o c t a spp.  25  XIII.  XIV.  of  Microfilariae  Measurements of Canadian m i c r o f i l a r i a e s i m i l a r t o Mf. f l e x i v a q i n a l i s Morphometric d e s c r i p t i o n s of Males S p l e n d i d o f i l a r i a p e c t o r a l i s n. sp. and S. p a p i l l o c e r c a  26  of ,  .  Morphometric d e s c r i p t i o n s of Females of S p l e n d i d o f i l a r i a p e c t o r a l i s n. s p . and S. p a p i l l o c e r c a Morphometric  descriptions  and Mf.  40  42  of Mf.  papillocerca  47  Morphometric descriptions of three s i m i l a r m i c r o f i l a r i a e from Canadian grouse  52  M i c r o f i l a r i a l infections i n the blue grouse examined from Vancouver Island (56 a d u l t m a l e s , 29 a d u l t f e m a l e s , a n d 16 y e a r l i n g s )  114  Measurements of similar microfilariae from grouse and p a s s e r i n e s  121  Range larvae  XII.  Page  Morphometric d e s c r i p t i o n s of Males of Skr.jabinocta f l e x i v a q i n - a l i s and s i m i l a r species. . . . . . . . '  lagopodis VIII.  TABLES  of of  measurements S.  of  pectoralis  Measurements  of  Microfilaria  sp.  fourth-stage n.  sp.  third-stage B  .  .  .  .  .  .  .  larvae .  .  .  .  .  .  .  .  .  .  137  of .  .  .  .  .  .  .  Some d e v e l o p m e n t a l a n d d e f i n i t i v e features of 6 avian onchocercids C o l l e c t i o n of s i m u l i i d s from arboreal evening exposures of d i f f e r e n t duration a t t h e same s i t e i n t h r e e y e a r s . . . . .  149  156  .  .  .  .  168  xi LIST OF TABLES (continued) TABLE XV,  XVI.  TITLE C o l l e c t i o n of C u l i c o i d e s from a r b o r e a l evening exposures of d i f f e r e n t d u r a t i o n at the same s i t e i n three years  Page  170  R e s u l t s of c o l l e c t i n g b i t i n g f l i e s from two grouse i n simultaneous a r b o r e a l and t e r r e s t r i a l evening exposures of same 174  XVII,  Comparison of s i m u l i i d c o l l e c t i o n s at the most p r o d u c t i v e s i t e and four other sites in i t s v i c i n i t y . . . . . . . . . .  177  xii LIST OF APPENDICES Page Appendix Table I. Occurrence of f i l a r i a l i n f e c t i o n s i n blue grouse and r u f f e d grouse from c o a s t a l B r i t i s h Columbia . . . . . . . . .  211  Appendix Table I I . Occurrence of f i l a r i a l i n f e c t i o n s i n g a l l i f o r m b i r d s of c e n t r a l B r i t i s h Columbia  214  Appendix Table I I I . Occurrence of f i l a r i a l i n f e c t i o n s i n t e t r a o n i d s of northern B r i t i s h Columbia and southern Alaska . . 220 Appendix Table IV. Adult n o n - g a l l i n a c e o u s b i r d s of B r i t i s h Columbia examined f o r m i c r o f i l a r a e m i a s 1959-62. . . . . . . . . . . . .  222  Appendix Table V. Q u a n t i t a t i v e d e s c r i p t i o n s of m i c r o f i l a r i a e from n o n - g a l l i n a c e o u s b i r d s i n B r i t i s h Columbia  224  Appendix VI. Q u a l i t a t i v e d e s c r i p t i o n s of microf i l a r i a e from n o n - g a l l i n a c e o u s b i r d s i n B r i t i s h Columbia  228  Appendix V I I . Geographic races of the four common t e t r a o n i d b i r d s of B r i t i s h Columbia  231  Appendix V I I I .  B i t i n g i n s e c t s f r o m the study 3 I 6 3 S » s  •  a  s  233  xiii LIST OF ILLUSTRATIONS PLATE I. II. III. IV.  .  TITLE  Page  Adults of S k r i a b i n o c t a f l e x i v a g i n a l i s . . . . . .  236  Adult Male S p l e n d i d o f i l a r i a  .  237  n. sp. .  238  n. sp. . . .  239  sp. A. . . . . . .  Adult Female S p l e n d i d o f i l a r i a p e c t o r a l i s Adult Male S. p e c t o r a l i s  n. sp.  Fourth-stage Larvae of S. p e c t o r a l i s V. VI.  Adult S p l e n d i d o f i l a ' r i a p a p i l l o c e r c a  240  M i c r o f i l a r i a sp. B and Mf. laoopodis  241  ACKNOWLEDGMENTS /The  author i s indebted  t o P r o f e s s o r James R.  Adams f o r h i s s u p e r v i s i o n , f o r h i s encouragement, h i s sound advice,  and f o r h i s a s s i s t a n c e  throughout the study. Mr.  i n many ways  Thanks are due Mr. D. Agur and  T. Barsby f o r p r o v i d i n g summer accommodation on the  study areas.  Members of the B r i t i s h Columbia F i s h and  Game Branch, p a r t i c u l a r l y Mr. P. M a r t i n , Mr. D. Robinson, and Mr. A. F r i s b y who generously  gave of t h e i r  c o n t r i b u t e d t o the study by o b t a i n i n g b i r d s and blood Dr.  smears.  time,  gallinaceous  The author wishes to thank  J . B. F o s t e r f o r c o l l e c t i n g  grouse from the Queen  C h a r l o t t e I s l a n d s , and Dr. R. B. Weeden f o r submitting filarioids  from Alaska  t e t r a o n i d s and f o r allowing- the  author t o examine a c o l l e c t i o n of blood  smears.  Thanks  are extended to the f o l l o w i n g i n d i v i d u a l s who made specimens of f i l a r i o i d s Dr.  available:  Dr. R. C. Anderson,  J . L. G r i t e s , Dr. R. L. Rausch, Dr. M. D. Sonin, and  Dr. M. J . Ulmer. The Mr.  a s s i s t a n c e of Mr. E. G. P l a t z e r and  P. T. K. Woo with  the f i e l d r e s e a r c h  i s gratefully  acknowledged.  P a r t i c u l a r thanks are extended to Mr. Woo  for  the author t o use some of h i s unpublished  permitting  data.  In the l a b o r a t o r y the t e c h n i c a l a s s i s t a n c e and  encouragement of Dr. Doris Jensen and Mrs. Z. J u r i c i c were much  appreciated.  XV Acknowledged provided a  B.  C.  by  a National  Sugar  Refining  The to  his  Committee  reviewing  the  go  J .  to  author  Dr.  F.  for  and  laboratory  careful  perusal  criticisms  their  and  to  Studentship  express  efforts  on  his  his  behalf  the  thesis.  Special  for  his  comments  on  with  several  To  thanks  thesis  suggestions.  Mrs.  and  phases  M. are her  B.  aid and  appreciation  of  sincere the  financial  Scholarship.  research.  of  the  Company wishes  assistance  is  Council  Bendell  Examiner,  thanks  Research  manuscript  and  External  his  for  with  the of  and  thanks thesis  the  Chitwood,  extended  for  for  field the her  thought-provoking  GENERAL  The (Weinland, f i l a r i a e , evolved  1858), or  and  tract  -  within  and the  With  the  brates  harbor  are  produce from  tebrate. the a  larvae  stage  The  There  which  vertebrate  infected  l)  of  horses,  of  which  fish,  worms,  parasitic  but  two  types.  they  digestive  adapted their  to  life  vertebrate  classes  of of  vertethese  In  place  one,  in  an  arthropod  the  f i l a r i a l  the  adult  females  inhabit  sites  eggs  have  access  to  the  exterior  ingested  by  the  intermediate  eggs  and  develop  and  life  embryonated  are  through  of  infecting  the  its  infection  either  or  arthropod  by  larvae  as  exemplifying  have  birds.  speaking,  broadly  the  majority  and  capable  i t  subcutaneously  which  feeds  this  P a r a f i l a r i a Yorke live  a l l  the  and  -  of  takes  acquires  the  cavities  of  highly  evolution  development  the  F i l a r i o i d s  species  serous  mammals  invertebrate,  proboscis skin.  and  they  is  the  successfully  eggs  leave  their  most  very  of  these  of  of  f i l a r i o i d s ,  the  become  thick-shelled  which  among  Filarioidea  as  roundworms  Larval  known  to  most  parasitize  cycles  are  course  f i l a r i a l  host  referred  specialized  exception  intermediate  superfamily  of  tissues  hosts.  helminths  the  habitat have  the  worms,  highly  During  the  of  generally  f i l a r i a l most  nematodes. forsaken  members  INTRODUCTION  and and  the  by  of  cycle  deposit  to  -host.  from  wounds  Maplestone,  host;  eating  escape  ver-  moults  d e f i n i t iv'e  around  type  two  of  the the  in  the  are 1926,  their  of  eggs  in  2 open haemorrhagic  l e s i o n s induced i n the s k i n .  are taken up by muscoid lesions 1909,  (37).  flies  (Haematobia)  T h e i r eggs  attracted  2) Species of D i p l o t r i a e n a • R a i l l i e t  of b i r d s , i n h a b i t  lungs,' are coughed  and Henry,  the a i r sacs; t h e i r eggs pass i n t o the  up, swallowed,  host's faeces (12).  to the  and reach the outside i n the  These eggs are i n g e s t e d by grasshoppers •  f e e d i n g on faeces or contaminated v e g e t a t i o n . In the other type -of c y c l e , the adult females produce p o o r l y d i f f e r e n t i a t e d but very a c t i v e embryos, known as m i c r o f i l a r i a e , which abound' i n the t i s s u e s by haematophagous arthropods.  and are i n g e s t e d  Some worms, such as Onchocerca  D i e s i n g , 1841, and S t e p h a h o f i l a r i a I h l e and Ihle-Landenberg, 1933,  live  subcutaneously,and t h e i r embryos accumulate i n the  s k i n , where they may ( S t e p h a n o f i l a r i a ) or may not (Onchocerca) be' a s s o c i a t e d w i t h l e s i o n s that a t t r a c t b i t i n g (43, 77). Most of the v i v i p a r o u s f i l a r i o i d s  vectors  (Loa S t i l e s , 1905,  Dipetalonema D i e s i n g , 1861, Wuchereria S i l v a Araujo, 1877, etc.) produce m i c r o f i l a r i a e which i n h a b i t the blood stream of t h e i r hosts; c o n c o m i t a n t l y , the adult worms occupy an extremely wide v a r i e t y of s i t e s deep w i t h i n the t i s s u e s . Of the.two dozen.genera filarioids  of m i c r o f i l a r i a - p r o d u c i n g  d e s c r i b e d from, b i r d s about 20 are placed by most  authors i n one subfamily - the S p l e n d i d o f i l a r i i n a e Chabaud and Choquet,  1953.  Almost h a l f of the 85-90 species of  s p l e n d i d o f i l a r i i n a e nematodes from b i r d s have been about e q u a l l y , to two- genera., Splendidof i l a r i a and C h a n d l e r e l l a Yorke  allocated,  S k r j a b i n , 1923,  and Maplestone, 1926 (10).  At l e a s t 20  3 new  s p e c i e s have been d e s c r i b e d i n these genera during the-past  decade,  and, c e r t a i n l y , many more have escaped d e t e c t i o n  cause of t h e i r small s i z e  be-  and o c c u l t s i t e s i n the hosts'*  t i s s u e s . . . Whereas adult v i v i p a r o u s f i l a r i o i d s collected infrequently, microfilariae veys of avian blood p a r a s i t e s .  are o f t e n noted i n sur-  General surveys of haematozoa  of b i r d s east of the Rocky Mountains Ontario  (18, 29), Maine  Georgia  (50), Iowa (3.3), Nebraska  where.  A few,  and Saunders  have been c a r r i e d out i n  (52, 55), D i s t r i c t  (31), and  (83)., else-  (60, 61) i n Georgia,  (63, 64) i n Mexico "have d e a l t infections,  of Columbia  (30), Texas  such as those of Robinson  with m i c r o f i l a r i a l  of b i r d s are  almost  and Robinson  exclusively  (60) found that  53 per cent of the avian s p e c i e s he examined harbored microfilariae.  General surveys of avian blood p a r a s i t e s from the  maritime r e g i o n of western North America have been c u r s o r y and few i n number (1, 35, 40, 42, 84), and although the present w r i t e r d e t e c t e d m i c r o f i l a r i a e  i n 38 per cent of 64  s p e c i e s of b i r d s from the western seaboard of Mexico l i s h e d data) the l i t e r a t u r e on the P a c i f i c  i s devoid of m i c r o f i l a r i a l  surveys  Coast.  Knowledge of f i l a r i a l  i n f e c t i o n s i n the g a l l i n a -  ceous b i r d s of extreme western North America fragmentary.  (unpub-  In 1944,  Beer  blue grouse i n Washington, adult f i l a r i o i d s  i s very  (17) surveyed the p a r a s i t e s of Idaho,  and Montana, but found no  and d i d not examine the blood.  In B r i t i s h  4 Columbia the presence of f i l a r i a l i n f e c t i o n s i n g a l l i n a c e o u s b i r d s was f i r s t  noted by Fowle  that 40 per cent of adult blue  (35) who r e p o r t e d , i n 1946, grouse from the Campbell  (Quinsam) r e g i o n of Vancouver I s l a n d harbored Schottelius  River  microfilariae.  (65) mentioned f i n d i n g m i c r o f i l a r i a e i n a few blue  grouse from c e n t r a l Washington i n 1951, but d i d not provide a recognizable corded  description.  microfilariae  In 1953, Adams and Bendell  i n 82 per cent  of adult blue grouse i n  the Quinsam r e g i o n , and from Alaska, Microfilaria and  lagopodis  (l) r e -  Babero (16) r e p o r t e d  "(Sambon, 1907)" i n w i l l o w  an u n i d e n t i f i e d subcutaneous f i l a r i o i d  ptarmigan  i n spruce  grouse.  About t h i s time, Game B i o l o g i s t s i n B r i t i s h Columbia became aware that u n i d e n t i f i e d f i l a r i a l worms occurred p e c t o r a l s k i n of blue grouse near Kamloops. l i s h e d r e c o r d of an i d e n t i f i e d  filarioid  under the  The only pub-  from b i r d s north of  Mexico i s t h a t of L o p h o r t o f i l a r i a c a l i f o r n i e n s i s Wehr and Herman, 1956, from q u a i l i n C a l i f o r n i a  (82) .  Prompted by the knowledge that f i l a r i a l i n f e c t i o n s were f r e q u e n t l y found i n blue grouse, and by the p a u c i t y of r e l e v a n t i n f o r m a t i o n , the w r i t e r s e t out to determine, the  species of f i l a r i a e  of t r a n s m i s s i o n . filarioids  Initially,  and second, t h e i r c y c l e  a t t e n t i o n was focused  of blue grouse alone, because  of the province g a I'll form,  i n t h i s host,  on the  l ) i n many regions  t h i s i s by f a r the most abundant n a t i v e  2) r e s e a r c h  advanced, and  first,  i n t o i t s bionomics was already w e l l  3) accommodation f o r f i e l d  r e s e a r c h was a v a i l -  5  able i n regions where blue grouse were p l e n t i f u l harbor  and known to  infections. Since blue grouse  (Dendraqapus obscurus  r e s i d e n t i n most of the b i o t i c  (Say)) are  zones of the province  d i s t r i b u t i o n o f t e n overlaps that of spruce  grouse,  their  ruffed  grouse, s h a r p - t a i l e d grouse and the i n t r o d u c e d chukar tridge,  species f o r which knowledge of f i l a r i a l  the r e g i o n i s e n t i r e l y  par-  infections i n  l a c k i n g . ' Therefore, when c o n s i d e r a b l e  data had been acquired on the f i l a r i a s e s of blue grouse, the study was expanded t o i n c l u d e the f i l a r i a l galliform  i n f e c t i o n s of other  birds. The  overall  aim of t h i s d i s s e r t a t i o n i s to provide  a comprehensive p i c t u r e of f i l a r i a l  i n f e c t i o n s i n the  t e t r a o n i d b i r d s of B r i t i s h Columbia and, to a l i m i t e d Alaska,  and the means by which these hosts  tions.  The t h e s i s i s presented  extent,  acquire t h e i r  i n two s e c t i o n s .  infec-  Part One  deals with the morphology and taxonomic p o s i t i o n of the filarioid  nematodes encountered i n t h i s study;  and Part Two  c o n t r i b u t e s new knowledge on the b i o l o g y of these a c o n s i d e r a t i o n of the f a c t o r s i n f l u e n c i n g t h e i r and d i s t r i b u t i o n i n the grouse of B r i t i s h  f i l a r i a e and transmission  Columbia.  6  PART  ONE.  MORPHOLOGY  AND  TAXONOMY  INTRODUCTION  The  writer  British  Columbia  species  (two  in  microfilariae tailed,  and  of  w i l l  f i l a r i a e  the  species  tailed, of  the  n.  comb.  Only  one  of  these  sites  (Syn.  to  described  single  grouse; as  male the  from  that  parasite  Columbia  grouse  that  and  Ohio  in and  (Jones,  Jones,  1961)  i n is  1961 1962  The  provisionally crows  most  1961)  Sultana,  Ohio.  of  blunt-  f i l a r i o i d  flexivaqinalis  in  two  occurs  This  crows  micro-  produces  flexivaqinalis  from  sharp-  Detailed  thinks  (Jones,  and  and  and  British  four  sites)  adults  worms  deciding  writer  species  species  the  sp.  A.  w i l l  be  of  to  be  described  circumoesophageal  designates  Splendidofilaria  third  for  Columbia  second  Reasons the  (45)  of  sheathed;  cavity.  Skriabinocta  reasons  he  This  body  deeper  of  should  writer the be  conspecific. The  a  why  flexivaqinalis  B r i t i s h  considered  the  originally  his  these  throughout  Splendidof i l a r i a  (78))  discuss  in  in  species.  f i l a r i a l  microfilariae.  similar  from  show  science.  Chandlerella  worms  w i l l  four  birds  f i l a r i o i d s  two  for  to  (45);  w i l l  presented  species  tetraonid  (blunt-tailed,  representing be  the  adult  sites,  new  various  that  harbor  are  sheathed  extremely  types  writer  tetraonid  habits  Alaska  two  descriptions  the  found  subcutaneous  unsheathed)  and  has  adult  this  new  is  fascia  based of  blue  species•temporarily  advanced.for  regarding  f i l a r i o i d  be  to  a  on  as  described,  new and  7 as an i n d i c a t i o n of i t s s i t e Splendidofilaria  i n the host, the name  p e c t o r a l i s n. sp. w i l l be proposed f o r i t .  Although the a d u l t s of S. p e c t o r a l i s n. sp. were found only i n grouse from the i n t e r i o r of B. C. and  from  Alaska, s h a r p - t a i l e d , unsheathed m i c r o f i l a r i a e - which are m o r p h o l o g i c a l l y i n d i s t i n g u i s h a b l e from Mf.  pectoralis  were very p r e v a l e n t i n blue grouse of Vancouver  Island.  Since i n t e n s i v e examination of a large number of grouse from t h i s r e g i o n f a i l e d  to l o c a t e the adult worms, the  w r i t e r w i l l designate t h i s c o a s t a l m i c r o f i l a r i a  as M i c r o f i l a r i a  sp. B. The f o u r t h species i n t e t r a o n i d s , i n t h i s case subcutaneously i n ptarmigan, i s S p l e n d i d o f i l a r i a (Lubimov,  1946)  Anderson  and Chabaud, 1959  Ularofilaria  p a p i l l o c e r c a Lubimov, 1946  papillocerca  (Lubimov,  1946)  This r a t h e r c o n t r o v e r s i a l  Spassky  (51); O r n i t h o f i l a r i a  and Sonin, 1957  f o r the f i r s t  A morphometric  (76)). from a  (51) - i s here r e p o r t e d  time, and the o r i g i n a l  inadequate d e s c r i p t i o n w i l l be supplemented mation.  (Syn.  species - f i r s t • d e s c r i b e d  g a l l i n a c e o u s host i n the U. S. S. R. from North America  (14)  papillocerca  w i t h new  infor-  d e s c r i p t i o n of i t s s h a r p - t a i l e d ,  unsheathed m i c r o f i l a r i a w i l l be presented; since blood smears are not a v a i l a b l e from the ptarmigan i n which adult p a p i l l o c e r c a were found, the m i c r o f i l a r i a  S.  i s described  from preserved specimens. S h a r p - t a i l e d , unsheathed m i c r o f i l a r i a e were present i n blood smears from ptarmigan not searched  8 f o r adult worms.  The w r i t e r w i l l  show why he thinks  these m i c r o f i l a r i a e are Mf. lagopodis  (Haaland, 1928) Brin'k-  mann, 1950, and w i l l point out and d i s c u s s that Mf. lagopodis  that  the p o s s i b i l i t y  i s a c t u a l l y the m i c r o f i l a r i a o f . S .  papillocerca. All  these species  belong to the subfamily  S p l e n d i d o f i l a r i i n a e Chabaud and Choquet, 1953, a group which has  given  considerable  (4,5,10,14) i n the l a s t the  difficulty  to taxonomists.  Anderson  few years has e x t e n s i v e l y  taxonomy of t h i s group, and has concluded  revised  (10) that the  l a c k of c o n s i s t e n t , major morphological d i f f e r e n c e s among the many s p e c i e s 1937,  previously  assigned  Gtinnert,  S p l e n d i d o f i l a r i a . L o p h o r t o f i l a r i a Wehr and Herman, 1956,  Chandlere11a. V a g r i f i l a r i a t h e i r being Skrjabin, The  to O r n i t h o f i l a r i a  placed  Augustine, 1937, e t c . ,  necessitates  i n only two genera, S p l e n d i d o f i l a r i a  1923 and C h a n d l e r e l l a Yorke and Maple:stone, 1926.  unwieldy s i z e of each of these c l o s e l y r e l a t e d genera  poses an o b s t a c l e  to the r e c o g n i t i o n of new s p e c i e s .  t h e l e s s , t h e i r number continues may be p o s s i b l e before  long  to i n c r e a s e .  Consequently, i t  to d i s c o v e r w i t h i n these genera  n a t u r a l groups which deserve independent generic haps on the b a s i s of c h a r a c t e r s p r e s e n t l y used f o r g e n e r i c The  writer w i l l  of- some of these c h a r a c t e r s  Never-  s t a t u s , per-  more s u b t l e than those  separation. consider and w i l l  the p o s s i b l e s i g n i f i c a n c e attempt to show t h a t more  sub-groups e x i s t n a t u r a l l y i n the genus C h a n d l e r e l l a  than are  9 now  recognized.  Chandlerella  The  removal of one  seems j u s t i f i e d  such sub-group from  at t h i s time, and  the w r i t e r  d i s c u s s h i s reasons f o r e l e v a t i n g t h i s group to the s t a t u s S k r i a b i n o c t a Chertkova,  With the  information  c r i p t i o n of a t h i r d , writer w i l l features  name Splendido-  sense proposed by Anderson  provided and  by two  observations  new  s p e c i e s , the  (.10)  (10). redes-  on s i m i l a r s p e c i e s ,  attempt to assess the v a l i d i t y of the  employed at present  generic  1946.  Throughout the t h e s i s the generic f i l a r i a w i l l be used i n the  will  the  morphological  to c h a r a c t e r i z e the  genus  Splendidofilaria. The  two  most recent  the  filarial  and  that of Sonin 1963,  nematodes, that of Chabaud and  advanced f o r c o n s i d e r i n g premature.  systems of c l a s s i f i c a t i o n  w i l l be the  compared and  Anderson  of  1959,  reasons w i l l  l a t t e r system i l l - f o u n d e d and  be  10  MATERIALS AND METHODS Hosts. writer  Gallinaceous  i n south-coastal  b i r d s were c o l l e c t e d by the  and s o u t h - c e n t r a l  as w e l l , the co-operation  British  Columbia;  of i n t e r e s t e d persons was e n l i s t e d  to o b t a i n  a few b i r d s from these areas and the Queen  Islands.  The carcases of f r e s h l y k i l l e d  or f r o z e n  Charlotte  gallinaceous  b i r d s were searched c a r e f u l l y f o r f i l a r i a l worms with the aid  of a d i s s e c t i n g microscope. Adult  worms.  L i v i n g f i l a r i a e were k i l l e d by  shaking i n c o l d 10% ethanol. 10%  ethanol  Specimens were preserved i n  5% g l y c e r i n e .  containing  For c l e a r i n g , the  a l c o h o l was allowed t o evaporate slowly were i n pure g l y c e r i n e .  Cleared  until  the worms  specimens were examined i n  temporary g l y c e r i n e mounts under supported c o v e r - s l i p s . quantitative  No  or q u a l i t a t i v e d i f f e r e n c e s were found between  f r e s h and thawed specimens of S p l e n d i d o f i l a r i a p e c t o r a l i s n.  sp.; the d e s c r i p t i o n of t h i s species  from f r o z e n and  blue grouse because these worms were most numerous  were i n u n i f o r m l y  nature of the adults  good c o n d i t i o n . of S k r i abinocta  specimens were u n s a t i s f a c t o r y d e s c r i p t i o n of t h i s species killed  i s based on specimens  Because of the f r a g i l e f l e x i v a q i n a l i s . thawed  f o r study; consequently, the  i s based on m a t e r i a l  from f r e s h l y  hosts. In a l l the d e s c r i p t i o n s  of-.'adult worms presented  i n t h i s s e c t i o n the average i s followed parentheses f o r each measurement.  by the range i n  11 In a d d i t i o n the  writer  tetraonid  to the specimens  r e c e i v e d subcutaneous birds For  collected  personally,  nematodes taken from  several  by s c i e n t i s t s i n Alaska. comparative purposes the w r i t e r fallisensis  examined  specimens  of S p l e n d i d o f i l a r i a  Anderson,  1961  (10)  Anderson,  1954  (3)) and S k r i abinocta chitwoodae  (Syn. O r n i t h o f i l a r i a  (Anderson,  1954)  fallisensis (Anderson,  1961)  n. comb. (Syn. C h a n d l e r e l l a chitwoodae  (11))  from the Ont a r i o Research Foundation, S k r i a b i n o c t a  flexivaginalis guiscali  from the Ohio State U n i v e r s i t y ,  (Linstow, 1904)  Linstow, 1904  (49); S p l e n d i d o f i l a r i a g u i s c a l i  Splendidofilaria  of the U. S. S. R.,  Ornithofilaria  Anderson,  1961  algonguinensis Anderson,  m i c r o f i l a r i a e were r a p i d l y and s t a i n e d .  and Mf. p e c t o r a l i s failed  (10)  1955  (Syn.  ( 4 ) ) , and an  collected  personally  Thin blood smears c o n t a i n i n g a i r - d r i e d , f i x e d i n absolute,  Somatic n u c l e i  from l i v i n g  of M i c r o f i l a r i a sp. B  or very f r e s h l y  k i l l e d hosts  to d i f f e r e n t i a t e i n Giemsa's s t a i n but were r e v e a l e d  s a t i s f a c t o r i l y by Wright's s t a i n or D e l a f i e l d ' s Dilute  algon-  birds.  Microfilariae.  methanol  1904)  University,  of S p l e n d i d o f i l a r i a  undescribed s p e c i e s of S p l e n d i d o f i l a r i a from Mexican  (Linstow,  from the Academy of Sciences  and specimens  (Anderson, 1955)  guiscali  (53)) from Iowa State  papillocerca  1961  Chandlere11a  n. comb. (Syn. F i l a r i a  Odetoyinbo and Ulmer, 1960  guinensis  Anderson,  haematoxylin.  Giemsa's s t a i n gave good r e s u l t s f o r M_f. f l e x i v a g i n a l i s .  and 'for.. Mf.  sp. B and Mf. p e c t o r a l i s  from s t a l e  or f r o z e n blood.  12 Microfilariae allowed  i n t h i c k smears or i n t h i n smears  to dry slowly were i n v a r i a b l y h i g h l y c o n t r a c t e d and  overstained;  consequently,  f o r morphometric study,  i n the t h i n n e r p o r t i o n s of r a p i d l y a i r - d r i e d  only  those  smears were used.  In a l l d e s c r i p t i o n s of m i c r o f i l a r i a e presented i n this  s e c t i o n the average length i s followed by the range i n  parentheses. so-called  The d i s t a n c e s  from the c e p h a l i c apex to the  " f i x e d p o i n t s " are expressed  as percentages of t o t a l  body l e n g t h ; the average i s followed by the range i n parentheses f o r each d i s t a n c e . A l l m i c r o f i l a r i a e d e s c r i b e d h e r e i n from blood f i l m s of western t e t r a o n i d s were i n smears made by the w r i t e r , with the exception  of Mf. laoopodis which the w r i t e r found i n  smears made by Dr. R. B. Weeden from ptarmigan.  On request,  members of the P r o v i n c i a l Department of R e c r e a t i o n  and Con-  s e r v a t i o n made blood smears from g a l l i n a c e o u s b i r d s at GameChecking S t a t i o n s i n the i n t e r i o r of B r i t i s h Columbia. smears c o n t r i b u t e d i n f o r m a t i o n on m i c r o f i l a - r i a l f o r a g r e a t e r number of host writer  alone  could have  occurrence  i n d i v i d u a l s and species than the  collected.  Smears of r u f f e d grouse blood c o n t a i n i n g aria  fallisi  These  Microfil-  Brinkmann, 1950, ( i n c l u d i n g the type i n d i v i d u a l s )  were borrowed from the O n t a r i o  Research Foundation i n order to  compare t h i s form with the s h a r p - t a i l e d m i c r o f i l a r i a e  of l o c a l  grouse. The  scientific  and common names used f o r l o c a l  t e t r a o n i d b i r d s i n the D e s c r i p t i v e S e c t i o n are as f o l l o w s :  Dendraaapus obscurus  (Say) - blue grouse  Canachites canadensis (L.) - spruce grouse Bonasa umbellus  (L.) - r u f f e d  grouse  Laqopus laqopus  (L.) - w i l l o w ptarmigan  Laoopus mutus (Montin) - rock ptarmigan L.  leucurus (Richardson) - w h i t e - t a i l e d  Pedioecetes p h a s i a n e l l u s  ptarmigan  (L.) - s h a r p - t a i l e d  grouse  14 RESULTS AND  DISCUSSION  I. DESCRIPTIVE .SECTION. A.  Filarioids  of C r y p t i c S i t e s  1. S k r i a b i n o c t a f l e x i v a q i n a l i s  (Jones,  a. D e s c r i p t i o n ( P l a t e I, f i g s . General:  Filarioidea,  S p l e n d i d o f i l a r i i n a e , S k r i abinocta elongate, usually very  n. comb,  1-7)  Onchocercidae, Chertkova, 1946.  Narrow,  d e l i c a t e worms, t a p e r i n g towards e x t r e m i t i e s ,  loosely coiled.  thin  1961)  (1-2  C u t i c l e l a c k i n g bosses or  | i t h i c k near mid-body).  Head sometimes set o f f  from body by s l i g h t c o n s t r i c t i o n at base.  O r a l opening  rounded by f o u r submedian p a i r s of p a p i l l a e . of i n n e r 4 p a p i l l a e .  striations,  Amphids at  surlevel  Oesophagus broad, d i v i d e d e x t e r n a l l y  i n t o r a t h e r short a n t e r i o r muscular p o r t i o n and  longer,  posterior, glandular  portion.  sometimes expanded.  J u n c t i o n of oesophagus with i n t e s t i n e  distinct,  a t r i c u s p i d valve  broad, t h i n - w a l l e d , No  t e r m i n a l caudal  Apex of muscular oesophagus  present..  I n t e s t i n e moderately  sometimes very broadly protuberances on e i t h e r  Male (see Table  dilated anteriorly. sex.  I f o r morphometric d e s c r i p t i o n ) :  P o s t e r i o r p o r t i o n of body u s u a l l y t i g h t l y c o i l e d . d i g i t i f o r m , with rounded, o f t e n knob-like cal  l i p bearing  apex.  Tail Anterior cloa-  a median p a p i l l a - l i k e protuberance  hypodermal connection.  Postanal  p a p i l l a e low.,  most f r e q u e n t l y 5 p a i r s , but v a r y i n g  with  subventral,  from 3 to 7 on e i t h e r side  15 (e.g., 4:3,  6:5,  dissimilar,  arcuate, only moderately s c l e r o t i z e d .  spicule left  18,  5:7)•Spicules  attenuate  left  subequal,  slightly Right  (maximum width about 25 \x) than  15 \x) , more s t r o n g l y curved  i n blunt point.  .slightly in  4:5,  longer and broader  (about  ending  3:5,  in distal  third.,  L e f t s p i c u l e with s h a r p l y pointed,  apex.  (Of 20 males, r i g h t s p i c u l e longer  s p i c u l e longer i n one,  s p i c u l e s equal i n  one.)  Heads of s p i c u l e s s c a r c e l y d i f f e r e n t i a t e d from blades, .seldom demarcated by any  constriction.  Walls  of s p i c u l a r heads  s l i g h t l y roughened i n t e r n a l l y , but not a p p r e c i a b l y Cells  and 4 to 6 n u c l e i of s p i c u l a r e p i t h e l i u m  w i t h i n a sac  thickened.  contained  (30 by 18 |i) at j u n c t i o n of r e t r a c t o r muscle  and  spicule. Female:  L i p s of v u l v a very s l i g h t l y  A n t e r i o r r e g i o n of vagina and 40  salient.  (? vagina vera) about 700  u. long  u. wide, t h i c k - w a l l e d , expanding r a t h e r a b r u p t l y  p o s t e r i o r p o r t i o n (? vagina width of body f o r 1-2 Microfilariae  mm  u t e r i n a ) which occupies  i n vagina measure 150-170 \i.  on r i g h t  entire  before becoming o p i s t h o d e l p h i c .  narrow, t u b e l i k e , d e l i c a t e , u s u a l l y short minating  into  side of body.  Rectum very  (about 40  u.) , t e r -  (In some specimens, anus  appears as minute pore i n s l i g h t d e p r e s s i o n ;  i n others,  patency of anus not d i s c e r n i b l e . )  Body narrowed r a t h e r  a b r u p t l y i n v i c i n i t y of anus, t a i l  of most females t e r m i n a t i n g  in  s l i g h t l y expanded knob. Microfilaria:  Phasmids  subterminal.  (85 specimens i n Giemsa-stained  smears of blood from 5 f r e s h and 4 f r o z e n adult blue  thin  grouse).  16 Moderate-sized with rounded e x t r e m i t i e s .  Generally  curved, or f l e x e d , seldom c o i l e d or sinuous. d e l i c a t e , h y a l i n e sheath p r o j e c t i n g 5-10 Length 170.3  about 0.6  u. apart.  f r e e of n u c l e i to 2.6  Nerve r i n g 24.4  C u t i c l e bearing very d e l i c a t e  A n t e r i o r t i p of m i c r o f i l a r i a  (1.8-3.9)%, t o t a l  c e p h a l i c space ex-  Somatic n u c l e i i n 3 r a t h e r compact rows.  (21.6-26.9)%,  Excretory v e s i c l e  always v i s i b l e  of moderate s i z e 36.0  E x c r e t o r y c e l l 39.5  occupying diameter of m i c r o f i l a r i a , Inner body s h o r t , 4.0  large,  subrectangular i n outline.  (58.8-64.4) % , p o s t e r i o r ;  First rectal cell  76.3  (72.3-81.4)%, occupying diameter of m i c r o f i l a r i a ,  of m i c r o f i l a r i a .  dot  (3.0-6.2)% of length of m i c r o f i l a r i a ,  (61.5-72.2)%.  i n outline  gap.  as r e f r a c t i l e  (37.3-40.5)%,  n o n - s t a i n i n g ; a n t e r i o r margin at 63.4 margin at 67.4  as narrow  (33.6-40.0)% to i t s  E x c r e t o r y pore sometimes v i s i b l e  or i n d e n t a t i o n .  circular  each end.  (6.7-10.0)%, c o n t a i n i n g 4 (2-7) l a r g e , somewhat  triangular nuclei.  mid-point.  u. beyond  Maximum width, c l o s e to a n t e r i o r end, 5.0-5.6 | i ;  width of caudal t i p 2.8-4.2 \x.  tends to 8.7  Surrounded by  (139.0-228.3) u., (70 were w i t h i n the range  150-185 u.) .  striations  straight,  i t s nucleus  and s l i g h t l y broader than h a l f diameter  Rectal c e l l s  2 to 4 r a t h e r evenly spaced  between R]_ and anal v e s i c l e , but obscure.  Anal v e s i c l e  (86.5-91.4)%, u s u a l l y s m a l l ; anal pore not observed.  89.5  Fifteen  to 20 n u c l e i present p o s t e r i o r to anal v e s i c l e , the l a s t 4 or 5 grouped near t i p , and f i n a l of apex.  2 c l o s e l y appressed to contour  17  Fourteen l i v i n g Mf. death of host, measured 193 b.  flexivaqinalis.  (178-209) | i .  Summary of.. occurrence i n B r i t i s h  *Hosts:  12 hours a f t e r  Columbia  Dendragapus obscurus f u l i q i n o s u s (Ridgway) D.  o.. s i t k e n s i s Swarth  D.  o. p a l l i d u s Swarth  D.  o,. r i c h a r d s o n i  (Douglas) - Mf.  only  Bonasa umbellus a f f i n i s A l d r i c h and  Friedman  B. u_. brunnescens Conover Canachites canadensis f r a n k l i n i Site:  (Douglas) - Mf.  In t h i n - w a l l e d mesenteric or p e r i t o n e a l throughout the body c a v i t y , u s u a l l y with the d o r s a l a o r t a , c o e l i a c , or mesenteric a r t e r y ; or w i t h i n the  Locality: c.  Coastal  and  c e n t r a l regions  I and and  II) have been placed  These are:  C.  (11)  associated superior  adrenals.  of the  species  province.  flexivaqinalis  which c l o s e l y resemble the w r i t e r ' s  Maplestone, 1926  1961  cysts  Discussion In a d d i t i o n to the  others  only  since Anderson's 1961  Skriabinocta  chitwoodae Anderson, 1961  H i b l e r , 1964  (44).  (Tables  i n the genus C h a n d l e r e l l a Yorke  C h a n d l e r e l l a petrowi (Syn.  specimens  three  r e v i s i o n (10).  (Chertkova, 1946)  Anderson,  petrowi Chertkova. 1946 (11),  and  (28)),  C. s t r i a t o s p i c u l a  These species w i l l be h e r e i n a f t e r r e -  * Those host species which y i e l d e d only the m i c r o f i l a r i a e of Sk. f l e x i v a q i n a l i s are so i n d i c a t e d .  18 ferred  to  in  Chertkova, confusion  combination  Justification Skri abinocta material (Jones,  Sk.  The  be  Jones'  (45)  qualitative  from  since  described blood  two  only  and  seems  Re i n t e r p r e t a t i o n  his  measurements  length  reveal  Jones'  evident  phology  microfilariae chitwoodae, possess or  crows  (or  shorter  present  from to 1  of  the  have  data  percentages  on  his  the  absence  natural versa).  glandular  of  and  the  of  papillae  rather  than  several  conversion  Sk.  (Table I I I ) . mor-  flexivaginalis  since  those  writer's  of  cross-infections,  "grouse  form"  opinion  that  the  writer's  oesophagus  and  longer  t a i l  Sk.  specimens  the  is  of  total  m i c r o f i l a r i a l  that  quanti-  paratypes.  (mean)  experimental  infections i t  of  of  material.  his  and  particularly  striatospicula.  In of  statement  par.atype  other  microfilariae  doubt  sheath,  on  a  longer  cephalic  vagina,  f a m i l i a r i t y with  no  any  misidentified  his  local  female  local  of  have  slightly  one  pairs  2  a  the  of  Sk.  vice  to  from  local  grouse  detect  lack  have  one.  discovery  to  s i m i l a r i t i e s  casts  are  and  to  avoid  flexivaginalis  from  and  to  to  The  II.  Sk.  females  unable  pairs  4  species  Section  male  Skriabinocta  Splendidofilaria)-.  these  to  name  binomials.,  for  oesophagus  structures. absolute  of in  microfilariae  smears,  "S."  differences of  in  "SJc."  the  was  observation  inaccurate,  Jones  but  examined and  generic  similarity  glandular  flexivaginalis, or  advanced  comb.,  writer  to  transfer  greatest n.  the  abbreviation the  w i l l  shorter  tative  is  for  1961)  t a i l .  the  shows  slightly  (abbreviated  1946 with  with  of  local  in the  females  19 do not j u s t i f y  separate  taxonomic s t a t u s f o r the worm from  grouse i n B r i t i s h Columbia, and that i t must be considered c o n s p e c i f i c with Sk. f l e x i v a q i n a l i s In 1903., E l l i o t t  (Jones, 1961)..  (32) d e s c r i b e d a sheathed  m i c r o f i l a r i a with an obtuse p o s t e r i o r extremity blood of a crow i n O n t a r i o . 1950  Microfilaria  clarkei  Brinkmann,  (20) from Bonasa umbellus -in O n t a r i o appears to be very  s i m i l a r , but i t i s r e p o r t e d to lack a sheath. failed both  from the  to l o c a t e the nerve r i n g or the anal v e s i c l e ,  seem to be present  (20, p. 13). sicle  although  i n h i s photograph of Mf. c l a r k e i  A l s o , he. was unable to f i n d the e x c r e t o r y ve-  (nor could he f i n d -it i n e i t h e r Mf. laoopodis or Mf.  fallisi). i n h i s type  Inasmuch as t h i s  structure i s c l e a r l y  visible  specimens of the l a t t e r , which the w r i t e r has  examined, i t i s concluded  that what he i d e n t i f i e d  e x c r e t o r y c e l l was, i n r e a l i t y The Table  Brinkmann  the e x c r e t o r y  as the  vesicle.  l i m i t e d measurements a v a i l a b l e , compared i n  IV i n d i c a t e pronounced . q u a n t i t a t i v e . s i m i l a r i t i e s among  these m i c r o f i l a r i a e .  This permits  the t e n t a t i v e s p e c u l a t i o n  that Sk. f l e x i v a q i n a l i s p a r a s i t i z e s t e t r a o n i d s as w e l l as c o r v i d s i n Eastern North America. The 1961)  n. comb., from a Java  striatospicula possess tively  species S k r i abinocta chitwoodae (-Anderson, sparrow ( l l ) , and Sk.  ( H i b l e r , 1964) n. comb., from magpies (44),  several quantitative s i m i l a r i t i e s small s i z e ,  short vagina,  such as compara-  and short male t a i l  which  20 d i s t i n g u i s h them from Sk. appears to be  separable  male, c o n s i d e r a b l y by possessing  flexivaginalis.  from Sk.  left  between 30  and 40  g r e a t e r diameter,  than r i g h t  unique among t h i s group of s p e c i e s ) . the i n n e r body of Sk.  H i b l e r (44)  more l i k e l y the c o r r e c t one.  other  c o i n c i d e with h i s  that the  l a t t e r length i s  As i s shown i n Table  i n n e r body of the m i c r o f i l a r i a e of Sk.  I I I the  chitwoodae and  Sk.  i s much s h o r t e r , a f u r t h e r p o i n t i n support  the d i s t i n c t n e s s of Sk:. petrowi  from the  broader, undivided  Chertkova, 1946,  three  from a d o v e ( 2 8 ) , i s  s i m i l a r species by  oesophagus, and  r a t h e r than a s s o c i a t e d with others.  of  striatospicula.  " i n t e r n a l media of the eye  as are the  ranged  Since the p o s i t i o n s of  d e s c r i p t i o n , the w r i t e r concludes  separable  s t a t e d that  u. i n l e n g t h ; however, h i s i l l u s t r a t i o n shows  fixed points in his figured m i c r o f i l a r i a  Sk.  and  s p i c u l e ( i n which i t i s  striatospicula microfilariae  an i n n e r body about 15 u- long.  flexivaginalis  chitwoodae  s t r i a t o s p i c u l a by i t s longer  longer female t a i l ,  a longer  Sk.  ;  1) i t s somewhat  2) i t s l o c a t i o n - i n the  adherent to v a s c u l a r membrane" blood v e s s e l s of the body c a v i t y ,  Table I.  Source of Data No. of Specimens Length  (mm)  Morphometric d e s c r i p t i o n s of Males of S k r i abinocta f l e x i v a q i n a l i s and s i m i l a r s p e c i e s . Measurements i n microns unless s t a t e d otherwise.  petrowi  chitwoodae  (28)  (11)  2  5  11,13.8  To Nerve Ring Length of T a i l  68  7.4-9.7  flexivaqinalis (writer;  grouse)  20 12.6(10.5-15.0)  flexivaqinalis (45;  crow)  35 .12.0(8.0-14.0)  84-111  140(121-160)  .136(124-140)  42*-68  89(66-114)  88(70-105)  Width of. Head.  33(30-37)  (27,31)*  W,  66(62-74)  (46,61)*  at Nerve Ring  Width at Anus  55  Maximum Width  150  L. Muse. Oesoph.  52(46-55) 90-99  126(110-160)  92-158  157(117-191)  striatospicula (44) 10 5(4-7) 99(90-135) 61(55-70)  48(28-53) (64,99)*  60(46-77)  137(129-159)  110(100-127)  Min.  W. Muse. Oesoph.  22(16-27)  18*  18  Max.  W. Muse. Oesoph.  28(25-34)  22*  22  (concluded next page)  ro  (Table  petrowi  W. Gland. Oes.  Max. W. Gland. Oes. L.  chitwoodae  0.440-1.38  L. Gland. Oes..(mm) Min.  I - concluded)  T o t a l Oes.  Long, Broad  (mm)  Spicule  L. of Long S p i c u l e L. of Short  Spicule  Postanal Papillae  64  flexivaginalis  flexivaginalis  (grouse)  (crow)  716(554-847)  743(533-911) (21,25)*  20  46(37-55)  (31,33)*  23  0.61-1.49  0.873(0.7201.03)  0.982(0.6601.05)  R  1R:4L  18R:IL  R  126  110(82-128)  7 pr.  77(67-90) 4-5 p r .  670(440-770)  35(30-47)  1.075  90  striatospicula  120(97-142)  0.780(ca.0.50.9) . R  105(85-126)  93(80-97)  99(74-114)  82(67-95)  65(50-68)  3-7 p r .  4-5 p r .  5-6 pr.  * From male paratypes examined by w r i t e r .  ro  Table I I . Morphometric d e s c r i p t i o n s of Females of S k r i abinocta f l e x i v a q i n a l i s and s i m i l a r s p e c i e s . Measurements i n microns unless stated otherwise.  petrowi Source of Data No. of Specimens Length  (mm)  (28) 1 27.5  To Nerve Ring To Vulva L. Vagina  322 (mm)  Length of T a i l  chitwoodae (11)  flexivaqinalis (writer;  grouse)  20  2  flexivaqinalis (45;  crow) 15  25.0(20.0-28.0)  154(123-209)  139(119-180)  141(98-184)  350,516  482(525-603)  466(350-575)  375(350-510)  0.750,0.330  2.0(1.0-3.1)  3.0(2.6-3.5)  150,200  237(141-394)  125(98-145)  W.  95(81-115)  67*  113(94-142)  147(98-182)  at Nerve Ring  280  14(13-18)  160,158  33*  Maximum Width  10  30.5(22.2-38.1)  36(28-41)  111  (44)  14.2,18.3  Width of Head  Width at Vulva  striatospicula  180,170  210(172-286)  0.705(0.650-0.8) 68(60-75)  116(96-128)  (concluded next page)  ro  (Table  petrowi  I I - concluded)  chitwoodae  153,142  L. Muse. Oesoph.  flexivaginalis  flexivaginalis  (grouse)  (crow)  192(145-248)  144(121-205)  striatospicula  160(135-184)  Min. W. Muse. Oes.  26(21-31)  18  Max. W. Muse. Oes.  32(28-36)  23  L. Gland. Oes.  (mm)  0.557,0.613  L. T o t a l Oes.  (mm)  1.24(0.9861.58)  41(31-46.)  Min. W. Gland. Oes. Max. W. Gland. Oes.  0.806(0.6500.904)  98 1.462  0.710,0.755  * From female paratype examined by w r i t e r .  0.975(0.4501. 10) 18  53(47-60)  42*  1.00(0.7951.12)  1.35(1.131.70)  23 1.09(ca.0.6-1.3)  Table  I I I . Morphometric d e s c r i p t i o n s of M i c r o f i l a r i a e of S k r i abinocta spp.  chitwoodae Anderson ( l l )  Source of Data  f lexivaainalis Writer  (grouse)  flexivaqinalis  striatospicula  Jones (45; crow)  H i b l e r (44)  No. o f Specimens  10  85  (?)  25  Sheath  +  +  0  +  Length (u.)  198(179-208)  170(139-228)  150(142-175)  Maximum Width (u.)  approx. 5  approx. 6  approx. 5  To Nerve Ring  (%)  24(21-28)  24(22-27)  -  Excr. V e s i c l e  {%)  35(34-38)  36(34-40)  (?45)  38(36-40)  39(37-41)  -  62(61-66)  63(59-67)  66(65-68)  67(62-72)  4  4(3.0-6.2)  (short)  74(73-76)  76(72-81)  (75)  87(86-89)  89(86-91)  Excr. C e l l Start End  {%)  of IB  of IB  {%)  {%)  L. of Inner Body 1st R e c t a l C e l l Anal V e s i c l e  * From vagina  {%)  {%) {%)  of preserved  (?65)  a  154(138-181) 6 23 ( 20 - 25) 34(30-38)  a  38(32-42)  (?64)  b  -  a  b  (23)  (58)  b  (67)  b  a  (9)  b  72(63-78) 83(73-88)  females.  a C a l c u l a t e d from the published  data.  b C a l c u l a t e d from the published  illustration.  ro  26  Table IV.  Measurements of Canadian m i c r o f i l a r i a e s i m i l a r to Mf. f l e x i v a q i n a l i s .  ' • . Mf. flexivaqinalis Source.of Data Locality  (this Br.  Host  study)  Columbia  Mf • clarkei (20) Ontario  , Elliott's Microfilaria (32) Ontario Crow  Grouse  Grouse  Length (u.)  170  178  173  Max.  5.6 .  5.7 .  5.8  Width (u.)  Cephalic Space {%)  5  4  36  37  37-40  {%)  63  62  ca.60  End of Inner Body'{%)  67  68  ca.70  Excretory V e s i c l e {%) S t a r t of Inner Body  Length of Inner Body  {%)  4(3.0-6.2)  6(3.6-8.8)  5  -  27 2.  S p l e n d i d o f i l a r i a sp. A a.  Description  (Plate  II, figs.  1-7)  F i l a r i o i d e a , Onchocercidae, S p l e n d i d o f i l a r i i n a e , Splendidofilaria Skrjabin, unless s t a t e d 48,. d o r s o v e n t r a l  1923.  otherwise.)  (One male; measurements i n  Width of head ( l a t e r o l a t e r a l  32) ; width of body at nerve r i n g 63; maximum  width 132; width at anus 48. •middle of nerve r i n g 176.  Distance from c e p h a l i c  Length of oesophagus 505; minimum  width of oesophagus 6, maximum width 9. 25.  Length of r i g h t s p i c u l e 78; length  Length of t a i l  38.  T o t a l length  Width of i n t e s t i n e of l e f t  s p i c u l e 84.  of worm 8.11 mm.  Head of worm compressed d o r s o v e n t r a l l y , ovoid  Two p a i r s of moderately large  papillae situated dorsoventrally);  at the "corners"  anterior papillae.  line,  plane; amphids at l e v e l of the more  C u t i c l e t h i c k , comprising .an outer smooth  inner  l a y e r with r a t h e r r e g u l a r  apart  bearing very f i n e transverse  and an  deep annulae about 10-15 us t r i a t i o n s averaging about  10 per annulus.  A papilla-like  precloacal  Right s p i c u l e with a short  structure  i s present on the head  at an angle to the s h a f t ; blade of s p i c u l e  dilated  side  considerably  l a y e r , up to about 6 u- t h i c k , but v a r i a b l e ;  lip.  (viewed  viewed a p i c a l l y the 2 p a p i l l a e of each  to the m i d - f r o n t a l  hyaline  cephalic  of the r e c t a n g l e  almost i n the same a n t e r i o r - p o s t e r i o r  dorsal  set  assuming  shape i n a p i c a l view and subreetangular shape i n dorso-  v e n t r a l view.  are  apex to  (6 |i long)  slightly  ( t o 10 u.) , then narrowed; a p i c a l one-quarter  slightly  28 curved ventrad, point. half  Left  slightly  spicule  thickened v e n t r a l l y ,  slightly  i t s l e n g t h , then  directed  tapering  to a sharp p o i n t ;  width  u,.  form  10  Caudal  protuberances b.  narrowed  papillae  9 | i , maximum  observed.  Terminal  (Ridgway)  papilli-  occurrence  Dendraqapus obscurus  fuliqinosus  Site:  In c o n n e c t i v e t i s s u e  o f neck, between t r a c h e a  c.  oesophagus.  Courtenay,  Vancouver  Island.  Comments . This s p e c i e s resembles  (Lubimov, -caperata  1946)  Anderson  Hibler.  1964  short  from  tail,  differing  that  and  from  characters,  S.  and  o n l y .S.  Chabaud,  (from the  p o s s e s s i n g a complex,  separated  than  and  Host:  Locality:  are  more  more v e n t r a d  o f i t s head  not  i n blunt .  present.  Summary.of  and  in  at a l i t t l e  slightly  width  ending  1959,  pulmonary  c a p e r a t a by  i t s lack  papillocerca  Splendidofilaria  i t seems r e a s o n a b l e very c l o s e l y  related.  (44)  I t can  o f magpie) be  s h o r t heads. i n the  last  A i s so s i m i l a r  t o suppose  S.  of caudal p a p i l l a e , i t s  as w e l l ,  sp.  and  arteries  annula'ted c u t i c l e .  i t s narrow s p i c u l e s w i t h S.  papillocerca  that  these  While  two  otherwise  two  species  29 B. F i l a r i o i d s 1.  of Subcutaneous  Sites  S p i e n d i d o f i l a r i a p e c t o r a l i s n. sp. a.  Description General:  (Plate I I I , f i g s .  1-6;  PI. I V , f i g s .  S p i e n d i d o f i l a r i a S k r j a b i n , 1923.  s l e n d e r worms, moderately  attenuated towards  1-3)  Long  extremities.  C e p h a l i c apex b l u n t , symmetrical, o f t e n s l i g h t l y broader than immediately p o s t - c e p h a l i c r e g i o n . slit,  o f t e n s l i g h t l y depressed.  O r a l opening a d o r s o v e n t r a l Buccal capsule absent.  Two  pairs cephalic papillae  and amphids arranged as f o l l o w s :  each l a t e r a l hemisphere  of apex, a large p r o j e c t i n g  papilla,  d o r s o l a t e r a l i n p o s i t i o n ; midway between i t and o r a l a small s e s s i l e p a p i l l a ;  amphids at l e v e l of large  j u s t v e n t r a l to m i d - f r o n t a l plane.  on  opening  papillae,  E x c r e t o r y pore and duct  not observed with c e r t a i n t y . ' Oesophagus moderately  long and  very s l e n d e r , comprising 3 continuous r e g i o n s as f o l l o w s : a n t e r i o r one-quarter to t w o - f i f t h s extremely narrow,  non-  g l a n d u l a r , but not o b v i o u s l y muscular; mid-region s e v e r a l as broad, i t s w a l l t h i c k e r  (4 |i) , g r a n u l a r , and  times  frequently  v a c u o l a t e d ; p o s t e r i o r one-quarter s i m i l a r to preceding r e g i o n but somewhat narrower,  l e s s g r a n u l a r , merging.with  without apparent valve or c o n s t r i c t i o n .  Intestine  30-46 u. i n diameter, i t s w a l l s about 6 u. t h i c k .  intestine narrow,  Cuticle  about  10 |i t h i c k over most of body, l a c k i n g bosses, and bearing f i n e transverse s u p e r f i c i a l  strations,  about  1 u. apart at mid-body.  Caudal apex of both sexes bearing a p a i r of prominent, v e n t r o l a t e r a l protuberances with hypodermal  rounded,  cores, sometimes  30 asymmetrically  developed.  Male:  (See  Table V f o r morphometric d e s c r i p t i o n )  T e s t i s u s u a l l y extending  to about 450  |i from c e p h a l i c apex.  P o s t e r i o r p o r t i o n of body c o n s i s t e n t l y curved i n shape of button hook, not c o i l e d . median l i p s .  C l o a c a l aperture bordered  Preanal l i p bears  invaded by hypodermis. small, v e n t r o l a t e r a l , subequal,  Caudal  single  l a r g e median  p a p i l l a e absent.  subterminal.  by p r o t r u d i n g protuberance  Phasmids very  S p i c u l e s moderately  stout,  s l i g h t l y d i s s i m i l a r ; head demarcated from blade  by  c o n s t r i c t i o n ; d i s t a l p o r t i o n of blade c u r v i n g a b r u p t l y ventrad. Right s p i c u l e  slightly  s h o r t e r than l e f t ,  ending  i n blunt,  s l i g h t l y expanded membranous l i p . L e f t s p i c u l e more s h a r p l y bent d i s t a l l y ,  ending  Female:  i n d u l l membranous p o i n t . (see Table VI f o r morphometric d e s c r i p t i o n )  L i p s of v u l v a w e l l developed, b r o a d l y rounded a n t e r i o r l i p . l y long, d i v i d i n g 15 |i broad; imparting behind  Vagina  i n t o two p a r a l l e l u t e r i .  s t r o n g l y r i d g e d appearance.  anus, sometimes to caudal apex.  anus patent. |i broad  \x long.  Walls of vagina not  about  contiguous,  Ovaries u s u a l l y  extending  Rectum d e l i c a t e , vacuo-  Perianal region s l i g h t l y elevated;  T a i l o f t e n c u r v i n g ventrad, ending b l u n t l y , about  at apex. Microfilaria:  (35 moribund m i c r o f i l a r i a e  made 12 to 24 hours post mortem host from 2 blue and grouse,  the  u s u a l l y s t r a i g h t , moderate-  apices of v a g i n a l e p i t h e l i a l c e l l s  l a t e d , about 100  40  very prominent, e s p e c i a l l y  and 35 m i c r o f i l a r i a e  1 ruffed  from thawed blood of 4 blue  f r o z e n about 5 hours post mortem; a l l grouse adults; m i c r o f i l a r i a e  i n smears  Giemsa-stained.)  with S.  grouse  pectoralis  Lacking sheath.  Cuticle  31 bearing f i n e t r a n s v e r s e s t r i a t i o n s n u c l e i i n 3-4 or  l o n g i t u d i n a l rows.  about 0.9  u. apart.  Somatic  A n t e r i o r end broadly  bearing n i p p l e - l i k e p r o j e c t i o n of o r a l apparatus.  rounded Anterior  extremity u s u a l l y s l i g h t l y broader than remainder of microfilaria.  Body t a p e r i n g r a t h e r evenly caudad from p o s t e r i o r  margin of i n n e r body. at  c e p h a l i c space 6.8  to  2.3  (1.5-5.2)%.  (164-236) u..  (5.9-8.4) u..  Maximum width,  A n t e r i o r end free of n u c l e i  C e p h a l i c space c o n t a i n i n g 4 or 5 large  g u l a r l y oval n u c l e i . Nerve r i n g  Length 188  Somatic n u c l e i beginning at 5.0  always v i s i b l e ,  at 16.4  (13.5-19.0)%.  irre-  (4.0-6.0)%.  Excretory  v e s i c l e up to 7 \x long, f r e q u e n t l y occupying l e s s than e n t i r e width of body, at 24.8  (20.2-29.0)% to mid-point.  pore sometimes v i s i b l e  as r e f r a c t i l e  vesicle.  Excretory c e l l  microfilarae in  beginning at 45.5  (26.1-33.2)%, u s u a l l y seen i n  Inner body conspicuous, seldom  eosinophilic,  (39.5-50.6)% and ending at 56.5  Length of i n n e r body 11.0  filaria;  spot about mid-point of  i n smears of thawed blood, not o f t e n seen i n those  f r e s h smears.  rectal cell  at 30.0  Excretory  at 64.0  (49.7-62.9)%.  (4.7-19.1)% of body l e n g t h .  (59.2-69.2)%, occupying diameter of micro-  nucleus of R-^ l a r g e .  c l o s e r to anal v e s i c l e  Rectal c e l l s  than to R^.  2 to 4 obscure,  Anal v e s i c l e  large,  gate, f r e q u e n t l y comprising s e v e r a l large h y a l i n e s t r u c t u r e s , 80.3 observed.  in  vacuolar  Frequently 4 or 5 s m a l l , i r r e g u l a r deeply s t a i n i n g  around or behind anal v e s i c l e .  a l o n g i t u d i n a l row of 3.  nuclei.  elon-  (75.8-83.3)% to mid-point; anal pore not  n u c l e i c l u s t e r e d around or behind anal v e s i c l e . clustered  First  Width of t a i l  Somatic  nuclei  Somatic n u c l e i ending  Terminal 2 t o 6 (i of t a i l  midway between anal v e s i c l e  lacking  and apex  32 5-7 (i; width of apex 1-2 | i ; t a i l  t i p u s u a l l y curved s l i g h t l y .  E i g h t moribund m i c r o f i l a r i a e  of S. p e c t o r a l i s  ina  blood-water mixture, 12 hours a f t e r death of host, measured: length 268 (238-279) \i; t o e x c r e t o r y pore 25.2 (24.5-25.8)%; t o s t a r t of i n n e r body 44,8 (44.2-45.9%; (53.8-57.5)%;  to end of i n n e r body 56.3  length of i n n e r body 11.5 (9.5-12.2)%; to anal  pore 78.4 (77.5-78.9)%.. Twenty m i c r o f i l a r i a e male S. p e c t o r a l i s  teased from the vagina of a f e -  preserved about 30 hours a f t e r death of host,  were s t a i n e d with Giemsa and mounted i n d i l u t e sealed c o v e r - s l i p .  These measured as f o l l o w s :  s t a i n under a length 201  (175-226) |_i; width of head 5 (4-6) (J,; maximum width 6 (5-7) to e x c r e t o r y v e s i c l e 47.9  u.;  24.2 (22.7-25.4)%; to s t a r t of inner body  (45.7-49.8)%; to end of i n n e r body 52.7 (49.7-54.4)%;  length of i n n e r body 4.8 (3.2-5.4)%; to f i r s t (60.0-62.7)%;  r e c t a l c e l l 61.2  to anal pore 78.1 (74.8-79.6)%; t a i l  t i p curved.  Apparently, most of the percentage d i s t a n c e s to f i x e d p o i n t s vary l i t t l e  between m i c r o f i l a r i a e  of d i f f e r e n t  lengths and treatments, except f o r the r e l a t i v e length of the i n ner  body which shows a marked r e d u c t i o n i n preserved specimens. b.  Hosts:  Summary of o c c u r r e n c e * Dendragapus obscurus pa 1 l i d u s  Swarth  Canachites canadensis a t r a t u s  Grinnell  C.  canadensis f r a n k l i n i (Douglas)  * Because of the s i m i l a r i t y between Mf. p e c t o r a l i s and Mf. sp. B, t h i s l i s t i n c l u d e s only those host species from which a d u l t s of S. p e c t o r a l i s were r e c o v e r e d .  33  Pedioecetes  p h a s i a n e l l u s caurus  Bonasa umbellus a f f i n i s Site:  Fried.  A i d . and  Fried.  Subcutaneous t i s s u e of p e c t o r a l r e g i o n , usually  about midway between crop and  t i p of sternum.  L o c a l i t y : Throughout B r i t i s h Columbia east of the Coast  Mountains,  from the Okanagan V a l l e y to the East Kootenay r e g i o n , and through the Cariboo  r e g i o n i n t o Alaska  McGrath) (see Appendix Tables c.  Peninsula,  II and I I I ) ,  Discussion B.abero (16) r e p o r t e d  ous  (Slana, Kenai  north  11  Filaria  sp. from the  subcutane-  t i s s u e over the p e c t o r a l muscles of seven Alaskan  grouse. Canachites  canadensis  (Linn;.)".  spruce  The w r i t e r has  examined  specimens of S. p e c t o r a l i s n. sp. from the same host i n Alaska, and p o s s i b l y t h i s i s the s p e c i e s to which Babero r e f e r r e d . S. p e c t o r a l i s i s . r e a d i l y . d i s t i n g u i s h e d m o r p h o l o g i c a l l y from the s p e c i e s of S p i e n d i d o f i l a r i a non-gallinaceous man,  1956)  (82). S.  hosts, and  Anderson, 1961,  (sensu Anderson, 1961)  from S. c a l i f o r n i e n s i s  of  (Wehr and  from the heart of C a l i f o r n i a  quail  It i s , however, r a t h e r s i m i l a r to S. p a p i l l o c e r c a and  tuvensis  filaria  (Spasskv  and Sonin,  t u v e n s i s Spasskv and  1957)  Sonin,  Anderson, 1961  separated  from S.  tuvensis  I t can  (76) by i t s lack of caudal  to  (= O r n i t h o -  1957), subcutaneous  s i t e s of g a l l i f o r m hosts i n the Soviet.Union.  Her-  para-  be papillae,  by i t s much more p o s t e r i o r v u l v a , b y the shape of the s p i c u l e s and  i n having  verse  a longer l e f t than r i g h t  i s true i n S. t u v e n s i s ) .  s p i c u l e (while  S. p e c t o r a l i s w i l l  with S. p a p i l l o c e r c a i n the next s e c t i o n .  be  the.concompared  34 2.  S p l e n d i d o f i l a r i a p a p i l l o c e r c a (Lubimov, 1946) a.  D e s c r i p t i v e note ( P l a t e One  the  surface  1-9)  male and two'female f i l a r i o i d s ,  c o l l e c t e d from  of the crop of ;a w h i t e - t a i l e d ptarmigan i n Alaska  •were r e c e i v e d Game.  V, f i g s .  from Dr. R. B. Weeden, Alaska  Dept. of F i s h and  These specimens were i d e n t i c a l t o those of S.  papillocerca  from T e t r a s t e s  bonasia r e c e i v e d  from the U.S.S.R.  A morphometric d e s c r i p t i o n of these specimens i s presented i n Tables V and VI, pp. .40-43, and on pp., 35-39 the d i s t i n c t i v e qualitative  features  are compared with those of S. p e c t o r a l i s .  An en face p r e p a r a t i o n bonasia r e v e a l e d  made from one of the females from T.  the f o l l o w i n g :  l a t e r a l l y , elongated e l l i p s e  the, c e p h a l i c apex i s a  i n the en face o u t l i n e with the 2.  p a i r s of p a p i l l a e and amphids s i t u a t e d almost completely ally;  later-  on each side the amphid i s s l i g h t l y v e n t r o l a t e r a l ; , very  near, the a n t e r i o r p a p i l l a , posterior papilla;  the p a p i l l a e are moderately prominent and  e q u a l l y w e l l developed.. same as t h a t o b t a i n i n g •b.  and d i r e c t l y a n t e r i o r to the  This c o n f i g u r a t i o n i s b a s i c a l l y the  i n S p l e n d i d o f i l a r i a , sp. A.  Summary of occurrence  Hosts and l o c a l i t i e s : Tetraogallus Tetrastes Tetrao  a l t a i c u s * Gebl .  bonasia L.  urogallus  Laqopus laqopus . : - ... Laqopus leucurus  i n U.S.S.R. ( A l t a i , 51) "  "  (Tuvinsk, 70.)  "  "  (  V. (L.)  "  (Richardson)"  11  11  _  70).:  (Kamchatka, Ukotsk, ... 74)  North America  (Alaska)  35 Site:  Subcutaneous, u s u a l l y  beside crop, or i n i n g u i n a l  region, c.  Discussion In 1879, Linstow (48) proposed the name  Filaria  u r o q a l l i f o r worms from the subcutaneous t i s s u e of Tetrao urogallus  i n Germany and B r i t a i n .  The lengths and widths of  males and females, r e s p e c t i v e l y , were given as: and  0.22 and 0=. 24 mm.  1/23  of body length;  cephalic  is  Other i n f o r m a t i o n i n c l u d e d : tail  oesophagus  of female rounded; head rounded,  apex somewhat f l a t t e n e d  prominences.  19 and 37 mm,  and surrounded by 6 f l a t  Probably, e i t h e r S. p a p i l l o c e r c a  or S.  tuvensis  F. urogal11. but any attempt to synoriymize one of the r e -  cent names with Linstow's would be groundless because of the inadequacy of h i s d e s c r i p t i o n .  The w r i t e r  c o n s i d e r s F.  u r o g a l l i Linstow, 1879 a nomen nudum because Linstow's t i o n does not embody s u f f i c i e n t I n f o r m a t i o n to permit  descriprecogni-  t i o n of the s p e c i e s he d e s c r i b e d . 3.  Comments on S. p a p i l l o c e r c a Lubimov's d e s c r i p t i o n  papillocerca and  indicates  and S. . p e c t o r a l i s  (51) of U l a r o f i l a r i a  many, s i m i l a r i t i e s between t h i s  species  S. p e c t o r a l i s n. sp.. and i s not d e t a i l e d enough that  points  of d i f f e r e n c e  can be e a s i l y determined.  The w r i t e r  examined 2 male and 5 female specimens from T e t r a s t e s  bonasia  ( c o l l e c t e d i n Tuva,. U. S; S.R. , 1957.) which had been determined as S. p a p i l l o c e r c a  by Dr. M... D. Sonin of the Academy of  Sciences of the U.S.S.R.  The q u a l i t a t i v e and q u a n t i t a t i v e  36 differences  which emerged from a comparison of the  specimens n e c e s s i t a t e s species.  t h e i r being considered  They have the  papilli'form structure  following  on p r e c l o a c a l  tuberances i n both sexes; no sharp-tailed tetraonid  c r i p t i o n s of S.  Tables V and  p e c t o r a l i s n.  most part,, q u a n t i t a t i v e nature; however, the twice as and  the  as  long.  tail  of female S.  The  following  papillocerca inner dex, 2-3  sp.  and  caudal pro-  rather two  long,  in  are  papillocerca. of a r a t h e r  papillocerca  p e c t o r a l i s . ' the  .For  the  subtle  i s more than  spicules  are  shorter,  i s only about o n e - f i f t h  striking qualitative  differences  groups of specimens.  c u t i c l e of both s p e c i e s i s t h i c k , that  of  comprising, from hyppdermis to e x t e r i o r ,  S.  a thick  l a y e r ; a t h i n l a y e r of s l i g h t l y d i f f e r e n t r e f r a c t i v e i n regularly  and  strongly  s t r i a t e d transversely  u. apart)-;, a l a y e r of i r r e g u l a r t r a n s v e r s e  resemble bosses a n t e r i o r l y ; and sometimes appears to be  sloughing o f f .  verse s t r i a t i o n s s l i g h t l y below the about 1 u. apart.  broadened from side head of S.  The  head of S.  to side  and  (striations  angulations which  a d e l i c a t e uneven l a y e r which  p e c t o r a l i s does.not comprise d i s t i n c t  and  S.  papillocerca  were noted on comparing the The  l i p ; terminal  VI present morphometric des-  differences  of S.  similarities:  a subcutaneous s i t e  vagina of S.  long as that  separate  caudal p a p i l l a e ; moderately  m i c r o f i l a r i a e ; (and  hosts).  qualitative  groups of  The  c u t i c l e of  l a y e r s , and  surface  the  are very  papillocerca  is  S. trans-  delicate noticeably  flattened dorsoventrally;  p e c t o r a l i s i s s l i g h t l y expanded s y m e t r i o a l l y .  the The  37  a n t e r i o r part of the body of S. p a p i l l o c e r c a t a p e r s r a t h e r g r a d u a l l y from the i n t e s t i n a l r e g i o n forward; t h i s r e g i o n of S. p e c t o r a l i s i s more markedly appearance.  the t a i l  double protuberance  narrowed, g i v i n g  a neck-like  of S . p a p i l l o c e r c a bears t e r m i n a l l y a  or a p a i r of s m a l l , acorn-shaped,  narrowly  separated protuberances; that of S. p e c t o r a l i s bears a pair, of l a r g e , rounded,  slightly  subterminal protuberances, sometimes  w i d e l y , sometimes narrowly separated. papillocerca pectoralis  The v u l v a r l i p s of S.  are weakly developed, not prominent:  are w e l l developed  a n t e r i o r lip...  those of S.  and p r o t r u d i n g , e s p e c i a l l y the  The apices of the v a g i n a l e p i t h e l i a l , c e l l s are  u s u a l l y contiguous i n S. p a p i l l o c e r c a : but are o f t e n not c o n t i guous i n S. p e c t o r a l i s . imparting to the vagina a s t r o n g l y r i d g e d appearance.  The a n t e r i o r l i p of the 'male c l o a c a  near i t s bas.e a. p a p i l l a - l i k e  s t r u c t u r e which i s r a t h e r small i n  S. p a p i l l o c e r c a . . and very prominent branous  bears  i n S. p e c t o r a l i s .  A mem-  apex i s absent from the s p i c u l e s of S. p a p i l l o c e r c a .  and present i n those of S. p e c t o r a l i s . broader than the l e f t as t h e : l e f t  The r i g h t  in. S. p a p i l l o c e r c a . and the same diameter  i n S. p e c t o r a l i s : i t s capituLum  parallel-sided  spicule i s  i s approximately  i n S. p a p i l l o c e r c a . and funnel-shaped i n S..  pectoralis. Anderson  (10) attached., c o n s i d e r a b l e importance to  the nature of the oesophagus i n h i s 1961 r e v i s i o n of Splendidofi1 aria  and r e l a t e d genera.  At. that time he i n c l u d e d  S. p a p i l l o c e r c a with s p e c i e s p o s s e s s i n g a "long, narrow, u n d i -  38 vided"  oesophagus,, "apparently  imperfectly  devoid  demarcated from i n t e s t i n e " .  (11), i n f l u e n c e d by the p a p i l l a - l i k e cloacal  of g l a n d u l a r  t i s s u e and  In a subsequent paper  s t r u c t u r e on the male pre-  l i p , and perhaps by Lubimov's f i g u r e . o f 0.098 mm f o r  oesophageal width, Anderson suggested that S. p a p i l l o c e r c a "needs restudy  as i t may belong i n C h a n d l e r e l l a " .  In the l i g h t  of Anderson's;statements and the o b s c u r i t y which surrounds S. papillocerca species  some comments on the oesophageal s t r u c t u r e of the  seem warranted.  The oesophagus of S. p a p i l l o c e r c a i s  narrow, as i s shown (Tables  V and VI) by. the measurements of  the w r i t e r and Sonin ( i n l i t t . ) , not broad, as Lubimov's f i g u r e of 0.098 mm  indicates.  p a p i l l o c e r c a two regions structure  Within  the oesophagus of S.  can be d i s c e r n e d ,  differing  slightly in  and i n diameter, and l a c k i n g abrupt t r a n s i t i o n .  The  a n t e r i o r p o r t i o n has a homogeneously h y a l i n e w a l l 5 to 6 u- t h i c k , and .a very  narrow lumen; the broader p o s t e r i o r p o r t i o n begins  j u s t behind the nerve r i n g and has a s l i g h t l y broader, e x t e r n a l l y more i r r e g u l a r w a l l , of somewhat v a r i a b l e t h i c k n e s s , but with numerous r e f r a c t i l e i n t e r n a l surface. and  a granular  cracks  or t h i c k e n i n g s  appearance, j u s t i n t e r n a l to the h y a l i n e wall,.  region represents  nature of t h i s  a shallow l a y e r of t i s s u e or the c u t i c u l a r  of the oesophagus.  r a t h e r asymmetrically, of two r e g i o n s ,  from i t s  This p o r t i o n has a somewhat broader lumen,  I t i s not c l e a r whether the apparent g r a n u l a r  lining  hyaline  The oesophagus j o i n s the i n t e s t i n e  but without a v a l v e .  Despite  t h i s oesophagus i s s u f f i c i e n t l y  ally., t o those of S. f a l l i s e n s i s  the evidence  similar, basic-  (Anderson, 1954) , S„  39  alqonquinensis similar  (Anderson, 1955), S. p e c t o r a l i s n. sp.  (undescribed)  s p e c i e s , which the w r i t e r has  that S. p a p i l l o c e r c a i s j u s t i f i a b l y in  i n c l u d e d with  examined,  these  species  Spiendidofilaria. It i s worth mentioning that the other  the p r e c l o a c a l protuberance, used by Anderson  character,  ( l l ) to suggest  a p o s s i b l e r e l a t i o n s h i p between S, p a p i l l o c e r c a and he placed i n C h a n d l e r e l l a . here considered (Sk. petrowi and  and  and  Sk.  S,. f a l l i s e n s i s .  to be  chitwoodae), i s present  2 species  i n S k r i abinocta  i n S.  as w e l l as i n S, p e c t o r a l i s n.  tuvensis sp.  Table V.  Source of Data  No. of Specimens  Morphometric descriptions- of Males of S p i e n d i d o f i l a r i a p e c t o r a l i s n. sp. and S. p a p i l l o c e r c a . Measurements i n mm.  S. p e c t o r a l i s  S. p a p i l l o c e r c a  Specimens from Br. C o l .  Specimens from U.S.S.R.*  20  Length  12.8(9.9-15.6)  To Nerve Ring  0.177(0.153-0.210)  Length of T a i l  0.109(0.082-0.140)  Lat.  0.074(0.071-0.078  W. Head  D-V Width Head W.  at Nerve Ring  11+,12.9 0.113  0.077,0.082 -  ,0.084  same  -  ,0.037  0.076(0.068-0.085)  -  ,0.095  Width at Anus  0.094(0.081-0.109)  Maximum Width  0.201(0.166-0.256)  L . Ant. Oesoph. Min.  -,  0.185,0.166 -  ,0.169  W. Ant. Oes,  0.009(0.006-0.012)  -  ,0.012  Max. W. Ant. Oes.  0.011(0.008-0.014)  -  ,0.014  Idem Specimens from Alaska  Idem Lubimov (51)  1  (?)2  10; 6  26-31  Idem Sonin**  20  0.086 0.055+  0.082  0.074  0.07(0.050.08)  0.098  0.148  0.196  0.08(0.050.09)  G.1.57 0.01-0.02  (concluded next page) o  (Table V - concluded)  . S. p e c t o r a l i s L. Post. Oesoph.  -  Min. W. Post. Oes.  0.016(0.010-0.019)  Max. W. Post. Oes.  0.029(0.021-0.034)  T o t a l Oesophagus  0.516(0.420-0.666)  L. of Rt. S p i c u l e  0.131(0.122-0.140)  Min,. W. Rt. S p i c . Max. W. Rt. S p i c . L. of L t . S p i c u l e  S. p a o i l l o c e r c a *  :  Idem  Idem  Idem**  0.332  -  -  0.020  -  -  - •  —  »  —  0.028  —  >  —  0.489  0.82  -  0 .093,lacking  0,095  0.102  -  0.016(0.012-0.021)  0 .012,  -  0.008  -  -  0.029(0.025-0.033)  • 0 .015,  -  0.014  -  -  0 . 110,lacking  0.106  0.087  -  . 0.139(0.131-0.145)  0.03  Min. W. L t . S p i c u l e  0.018(0.014-0.022)  0 .011,  -  0.008  -  -  Max. W. L t . S p i c u l e  0.028(0.023-0.031)  0 .012,  -  0.013  -  -  * Specimens from T e t r a s t e s bonasia r e c e i v e d from M. D. Sonin. ** Data from Sonin, .in l i t t .  Table VI.  Source of Data  No.  of Specimens  Morphometric d e s c r i p t i o n s of Females of S p l e n d i d o f i l a r i a p e c t o r a l i s n. sp. and S. p a p i l l o c e r c a . Measurements i n mm.  S_. p e c t o r a l i s  S. p a p i l l o c e r c a  Specimens from Br. C o l .  Specimens from U.S.S.R..*  Idem Specimens from Alaska  20 . 40.4(34.7-49.3)  37.3(28.7-42.9)  To Nerve Ring  0.191(0.169-0.243)  0.156(0.144-0.172)  To Vulva  0.622(0.420-1.057)  0.467(0.382-0.549)  L.  0.721(0.527-1.022)  1.52(1.30-2.030)  L. of T a i l  0.997(0.527-1.786)  0.178(0.119-0.262)  Lat. W. Head  0.095(0.078-0.113)  0.086(0.076-0.096)  same  0.050(0.043-0.058)  0.097(0.086-0.114)  0.108(0.092-0.117)  0.123,0.142  0.167(0.140-0.204)  0.136(0.107-0.150)  0.142,0.166  D-V Width Head W.  at Nerve Ring  Width at Vulva  Lubimov (51.) (?)2  Length  of Vagina  Idem  35.7,31+ -  Sonin**  20  48-50  ,0.138  0.358,0.418  0.470  2,09,0.95  2.040  0.283,0.105  0.306  -  Idem  0.15(0.090.21)  ,0.111  (concluded next page) 4^  to  (Table VI - concluded)  S. p e c t o r a l i s Maximum Width L. of Ant. Oes.  0.342(0.307-0.396) -  S. p a p i l l o c e r c a *  Idem  Idem  Idem**  0.300  -  0.132,0.172  -  -  0.253(0.215-0.320)  0.281, -  0.178(0.161-0.191)  Min. W. Ant. Oes.  0.006(0.005-0.010)  0.013(0.011-0.015)  0.007,0.014  -  Max. W. Ant. Oes.  0.011(0.005-0.015)  0.018(0.015-0,021)  0.017,0.014  -  -  0.339(0. 288-0.392)  0.392,0.418  -  -  :  L. Post. Oesoph.  -  0.03-0.04  Min. W. Post. Oes.  0.015(0.006-0.026)  0.026(0,023-0.030)  0.032,0.031  -  Max. W. Post. Oes.  0.026(0.018-0.035)  0.032(0,026-0.037)  0.037,0.034  0.098  0.05-0.06  T o t a l Oesophagus  0.759(0.493-0.875)  0.517(0.468-0.560)  0.524,0.590  0.570-0.64  0.5700.648  Pres. Mf ex ute.ro  0. 201(0.176.-0.226)  0.186(0.174-0.196)  0.190(0.1800.197)  * Specimens from T e t r a s t e s bonasia r e c e i v e d from M. D. Sonin. ** Data  from Sonin 1961 (71) and i n l i t t .  CO  44  C.  M i c r o f i l a r i a e of Unknown Adults 1.  Microfilaria a.  laqopodis  (Haaland,  1928)  Comparison with other d e s c r i p t i o n s (Plate VI, f i g .  6.)  A number of s p e c i e s of m i c r o f i l a r i a e , u n c o r r e l a t e d with  adult nematodes, has  p h a s i a n i d b i r d s (34, 55,  been d e s c r i b e d from t e t r a o n i d and 67, 80,  and o t h e r s ) .  Most of these  p o o r l y d e s c r i b e d and have been r e p o r t e d only once. tion is Microfilaria  laqopodis  which Brinkmann (19) concluded Sambon, 1907*  (Haaland, was  i n Alaska  tetrix  1950,  smithi  (63), Laqopus lagopus  i n Norway (75), and  in  from Laqopus  (16).  Brinkmann (19) presented what he considered to be Mf.  a d e t a i l e d d e s c r i p t i o n of  laqopodis.  s h a r p - t a i l e d , unsheathed m i c r o f i l a r i a e  The w r i t e r has  found  i n blood smears from  and L. mutus of extreme northern B r i t i s h Colum-  b i a which d i f f e r s l i g h t l y  from Mf.  but which c l o s e l y resemble Mf. mann.  Brinkmann,  Under these names, m i c r o f i l a r i a e have been repor-  Norway (19, 40), Lyrurus  Laqopus laqopus  excep-  i d e n t i c a l to F i l a r i a  ted from Laqopus s c o t i c u s i n B r i t a i n  laqopus  1928)  The  are  pectoralis  (and Mf_.  laqopodis as d e s c r i b e d by  Table VII provides morphometric d e s c r i p t i o n s of  laqopodis a f t e r Brinkmann (19),  sp. B) ,  2) m i c r o f i l a r i a e  Brinkl ) Mf.  from B.  C.  * S h i p l e y (68) noted i n 1909 that Sambon's name f o r t h i s m i c r o f i l a r i a , F i l a r i a s m i t h i . was preoccupied. Haaland (40) suggested the replacement name F i l a r i a l a q o p o d i s . and Brinkmann (19) p o i n t e d out that the name F i l a r i a was inapprop r i a t e , and proposed the combination M i c r o f i l a r i a l a q o p o d i s .  45 ptarmigan, and the preserved  3) Mf. Soviet  p a p i l l o c e r c a teased  and  Alaskan  from the vaginas  of  specimens of S., p a p i l l o c e r c a  available. The  most s a l i e n t  the w r i t e r ' s smears are  f e a t u r e s of the m i c r o f i l a r i a e i n  l) a s l i g h t l y i n f l a t e d cephalic region,  2) a moderately long c e p h a l i c space, ring the  and  4)  a long i n n e r body.  features reported  mann (19) single  3) a very d i s t i n c t  These observations  specimens was  to 8 | i " .  He  agree with  or i l l u s t r a t e d by other w r i t e r s .  found that i n h i s m i c r o f i l a r i a e  "the  nerve  Brink-  a n t e r i o r end  wider" than body diameter "being  of  found  up  d e s c r i b e d the i n n e r body as "some 22-30 u. i n  l e n g t h " , while  S/^rum (75)  t h i s s t r u c t u r e occupied  (quoted  by Brinkmann) s t a t e d that  about one-quarter of the t o t a l  length i n  1  h i s specimens from Lvrurus  tetrix.  Because of the s i m i l a r i t i e s  and  the  d i f f e r e n c e s between the w r i t e r ' s specimens and mann and  o t h e r s , i t i s probable  N e a r c t i c ptarmigan are Mf.  and Mf.  Brink-  laqopodis.  In the  light  of  the  identical microfilariae  t h i s determination  (Mf.  species  i s only t e n t a t i v e ,  Comparison with Mf. p a p i l l o c e r c a The  described,  of  sp. B) can be produced by d i f f e r e n t  of adult f i l a r i o i d s , b.  those  that the m i c r o f i l a r i a e i n  w r i t e r ' s d i s c o v e r y that v i r t u a l l y pectoralis  lack of obvious  microfilaria  as such.  of S. p a p i l l o c e r c a has  Unfortunately,  blood  never been  smears are not  avail-  able from the b i r d s from which adult S. p a p i l l o c e r c a were recovered.  Table VII i n c l u d e s the morphometric data  for  Mf.  46 papillocerca  teased from preserved females.  Mf. p e c t o r a l i s to those  that such m i c r o f i l a r i a e  I t was shown f o r  are somewhat d i s s i m i l a r  i n blood smears, p a r t i c u l a r l y i n the length of the  i n n e r body, which i s much l e s s i n preserved m a t e r i a l . the i n n e r body of preserved Mf. p a p i l l o c e r c a  Since  i s longer than  that of preserved Mf. p e c t o r a l i s . probably the i n n e r bodies of smeared m i c r o f i l a r i a e ship.  of these 2 species bear the same r e l a t i o n -  I f t h i s i s t r u e , then the i n n e r bodies of Mf. p a p i l l o -  cerca and Mf. lagopodis are s i m i l a r . similarities Mf. lagopodis microfilaria  T h i s , along with the other  of morphology, host, and d i s t r i b u t i o n suggests that (Haaland,  1928) Brinkmann, 1950 may w e l l be the  of S p l e n d i d o f i l a r i a  Anderson and. Chabaud, 1959.  papillocerca  (Lubimov, 1946)  Table V I I . Morphometric  d e s c r i p t i o n s of Mf. lagopodis and Mf. p a p i l l o c e r c a .  Mf. lagopodis on blood smears Host  L. lagopus  Source of Data  Norway (19)  No. of Specimens  L. lagopus B.C.(this  study)  21  Length (u-) To Nerve Ring  Mf. p a p i l l o c e r c a teased from p r e s e r v e d females  228(218-255) {%)  16,6(15,0-17.6) (?) 25.7(23.5-27.6)*  To Excr. Ves. {%)  215 ( 203 - 238)  T. bonasia  L. lagopus  U.S.S.R. ( t h i s study)  Alaska ( t h i s study)  16  10  186(174-196)  190(180-197)  -  15.7(14.5-17.0)  -  24.8(23.0-26.0)  24.6(23.4-27.7)  26.2(25.6-27.4)  To S t a r t of IB {%)  -  40.8(37.9-46.1)  46.8(44.3-48.8)  48.6(45.7-50.6)  To End of IB {%)  -  56.6(52.6-60.3)  52.8(51.8-53.6)  55.5(53.0-57.0)  15.8(13.7-21.6)  6.0(4.5-7.5)  6.9(6.0-8.8)  Length of IB {%) To 1  s t  Rectal C e l l  To Anal Ves. {%)  (?ca. {%)  10-13)  66.1(64.4-67.4)  66.2(62.9-71.2)  61.2(60.0-62.7)  77.7(72.1-85.2)  78.6(74.8-85.8)  77.0(75.2-79.2)  * Brinkmann's "excretory c e l l " .  79.0(77.5-81.4)  48 2.  Microfilaria a.  sp. B  Description  (Plate  VI,f i g .  l)  (Based on 40 w e l l extended specimens i n t h i n smears made up to 3 hours post mortem from 3 adult blue grouse, and 50 specimens  i n t h i n smears from 6 adult blue grouse and 1  adult r u f f e d grouse thawed s h o r t l y before smears were made; Giemsa-stained.) ... Moderately long, r a t h e r robust m i c r o f i l a r i a e , l a c k i n g a sheath, and w i t h f i n e t r a n s v e r s e  cuticular  stria-  t i o n s about 1 |i apart.  Length 214.6 (177.2-265.1) | i . Maximum  width 7.2 (5.6-7.8) u..  Region of c e p h a l i c  tapered to blunt  space  frequently  apex, but sometimes broadly rounded.  Cephalic  space c o n t a i n i n g 2-6 n u c l e i which extend forward to 2.3 (1.4-3.0)%; p o s t e r i o r  limit  of c e p h a l i c  space at 4.9 (3.7-6.7)%.  Somatic n u c l e i i n 3 or 4 l o n g i t u d i n a l rows. (15.3-20.6)%, o f t e n i n c o n s p i c u o u s . 26.2  Nerve r i n g  Excretory v e s i c l e  at 17.7  large,  (24.1-30.2)% to midpoint; e x c r e t o r y pore sometimes  visible  as minute p i n k - s t a i n e d dot.  E x c r e t o r y c e l l subrectan-  g u l a r , occupying width of body, but seldom s t a i n i n g w e l l , at 30.0  (27.0-33.8)%.  A n t e r i o r margin of i n n e r body at 48.1  (43.5-53.1); p o s t e r i o r margin at 57.8 (51.9-62.3)%. body of most specimens  f i l l e d with p i n k - s t a i n e d granules, and  occupying 9.7 (5.0-18.8)% of length of m i c r o f i l a r i a . rectal cell large  First  at 65.1 (60.4-69.2)%, i t s nucleus f r e q u e n t l y very  (extending almost across body); R2 seldom e v i d e n t ; R^  and R^ c l o s e together, near anal v e s i c l e . cell  Inner  s i m i l a r to R-^ v i s i b l e  Anal v e s i c l e  Sometimes a large  about 5 |i behind anal v e s i c l e .  at 81.7 (77.3-85.1)% t o mid-point, u s u a l l y ap-  49 pearing  as two tandem v a c u o l e s .  anal v e s i c l e , the t e r m i n a l irregular,  4 or 5 of which are i n a s l i g h t l y  l o n g i t u d i n a l row; t e r m i n a l  Width of body midway between anal Apex r a t h e r one  About 15 n u c l e i p o s t e r i o r to  sharply  pointed  4 u, l a c k i n g n u c l e i .  v e s i c l e and apex, about 4 | i .  and u s u a l l y curving  s l i g h t l y to  side. Four s l u g g i s h M i c r o f i l a r i a  sp. B i n whole grouse  blood 12 hours post mortem host ranged between 240 and 256 \x in  length. b.  Summary of occurrence*  Hosts:  Dendragapus obscurus f u l i q i n o s u s  (Ridgway)  D. o. s i t k e n s i s Swarth Bonasa umbellus brunnescens Conover Localities:  Vancouver I s l a n d , Queen C h a r l o t t e and  c.  south  coastal B r i t i s h  Islands,  Columbia.  Discussion A f t e r examining and measuring many hundred micro-  filariae  i n blood smears from grouse harboring  adult S.  p e c t o r a l i s and i n blood smears from Vancouver I s l a n d (Mf.  sp. B) the w r i t e r was f o r c e d  to conclude that  grouse  the minor  d i f f e r e n c e s between the 2 m i c r o f i l a r i a e were not of s e p a r a t i v e value.  This comparison was hampered by the f a c t that  smears c o n t a i n i n g  a l l blood  Mf. p e c t o r a l i s had been made e a r l y i n the  * The data i n Appendix Table I I suggest that the f o l l o w i n g grouse may be i n f e c t e d i n the i n t e r i o r of the p r o v i n c e : D, o. p a l l i d u s Swarth, and D. o. r i c h a r d s o n i (Douglas).  50 study and were t h i c k e r , had not been a i r - d r i e d  so r a p i d l y  and  were not so w e l l s t a i n e d as those made l a t e r on Vancouver Island. was  The  average  l e n g t h of Mf.. p e c t o r a l i s i n blood smears  l e s s than that of Mf.  sp. B.  However, i t i s probable  t h i s i s due, p a r t l y at l e a s t , to a l l o w i n g the smears with p e c t o r a l i s t o dry more slowly than those with Mf.  that Mf.  sp. B,  Despite d e t a i l e d examination of the carcases of 37 blue grouse  from c o a s t a l B r i t i s h Columbia,  i n f e c t e d with Mf.  a l l of which were  sp. B (many with high m i c r o f i l a r a e m i a s ) ,  the only f i l a r i o i d  found, other than Sk.. f l e x i v a g i n a l i s .  the s i n g l e male, d e s c r i b e d h e r e i n as S p l e n d i d o f i l a r i a Apparently, then, Mf.  sp. B i s not the m i c r o f i l a r i a  was  sp. A.  of S.  p e c t o r a l i s n. sp. or of S. p a p i l l o c e r c a . the a d u l t s of which are e a s i l y  l o c a t e d , and i n the absence  be concluded with c e r t a i n t y t h a t Mf. Splendidofilaria  sp. A.  sp. B i s the embryo of  Because of the extreme  between Mf. p e c t o r a l i s and Mf. decide which s p e c i e s was i n those grouse  of females i t cannot  similarity  sp. B i t i s not p o s s i b l e to  present (or whether both were present)  from the i n t e r i o r of the p r o v i n c e , i n f e c t e d  with s h a r p - t a i l e d m i c r o f i l a r i a e , but i n which S.  pectoralis  a d u l t s were not found, or which were not searched f o r adult filarioids  (see Appendix Table I I ) . Mf.  fallisi  Brinkmann, 1950  was  d e s c r i b e d from  smears of O n t a r i o r u f f e d grouse i n the c o l l e c t i o n of the O n t a r i o Research  Foundation  sonal communication,  1965)  (20).  Dr. R. C. Anderson  (per-  s t a t e s that the a d u l t s of Mf.  51 fallisi tions  have not  yet  of i n f e c t e d  interesting  been found, although numerous examina-  grouse have been c a r r i e d  to note (Table VIII) the  tween these two  m i c r o f i l a r i a e , the  escaped d e t e c t i o n .  Were i t not  of these i s d i s t i n g u i s h a b l e might be Mf.  sp.  out.*  marked s i m i l a r i t y  Mf.  fallisi  f o r the  from Mf.  ever, there i s no way  are  one  f a c t that  pectoralis  species.  of knowing how  As  many (?  microfilariae above three  Indeed, i t i s d i f f i c u l t of S.  (see  papillocerca  preceding s e c t i o n ) ,  information indicates microfilariae  are  (?Mf.  at  there that  i t i s , howsimilar) identical  to d i s t i n g u i s h  the  laqopodis) from  the  although the  rather conclusively  produced by  neither  conclusion  s p e c i e s of avian f i l a r i o i d s produce v i r t u a l l y microfilariae.  be-  a d u l t s of which have  some j u s t i f i c a t i o n f o r a t e n t a t i v e B and  It i s  that  available  these 4  l e a s t 3 s p e c i e s of  adult  females.  * The m i c r o f i l a r i a of A p r o c t e l l a s t o d d a r d i Cram, 1931, adults of which have been c o l l e c t e d from r u f f e d grouse i n Ontario (5), can be d i s t i n g u i s h e d e a s i l y from Mf. sp. B and Mf. fallisi.  Table V I I I .  Morphometric d e s c r i p t i o n s of three s i m i l a r m i c r o f i l a r i a e from Canadian grouse.  Mf. p e c t o r a l i s  Mf. f a l l i s i * 25  75  No. of Specimens  Mf. SD. B 90  Length (u.)  188(164-236)  194 ( 164-234)  215(177-265)  Max. Width (u.)  6.8(5.9-8.4)  5.9( 5.6-6.7)  7.2(5.6-7.8)  Ceph. Space  5.0(4.0-6.0)  5..K 3.7-6.2)  4.9(3.7-6.7)  {%) {%)  16.4(13.5-19.0)  18.7(  To Excr. Ves. {%)  24.8(20.2-29.0)  28.0( 26.2-30.0)  26.2.(24.1-30.2)  To Excr. C e l l  30.0(26.1-33.2)  31.0 ( 28.6-33.2)  30.0(27.0-33.8)  45.5(39.5-50.6)  49.3( 46.5-52.2)  48.1(43.5-53.1)  To Nerve Ring  {%)  To S t a r t of Inner Body  {%)  16.3-21.2)  17.7(15.3-20.6)  To End.of Inner Body  {%)  56.5(49.7-62.9)  57.0( 52.5-61.9)  57.8(51.9-62.3)  To F i r s t Rectal C e l l  {%)  64.0(59.2-69.2)  66.3( 63.6-70.3)  65.1(60.4-69.2)  80.3(75.8-83.3)  82.8(  81.7(77.3-85.1)  To Anal Ves. {%)  *  78.4-87.1)  From the w r i t e r ' s examination of Brinkmann's types.  ro  53 II. A.  TAXONOMIC SECTION  C h a n d l e r e l l a . S p i e n d i d o f i l a r i a . and Related Genera Numerous p r o p o s a l s have been made r e g a r d i n g the  d i s t i n c t n e s s of the genera C h a n d l e r e l l a Yorke 1926,  S p i e n d i d o f i l a r i a S k r j a b i n , 1923,  Gfinnert, 1937, mies.  1961.(10).  and O r n i t h o f i l a r i a  p r e f e r s . Anderson's  In the f o l l o w i n g pages a b r i e f  system.  synony-  to the b a s i c scheme of  summary w i l l ' be presented and the w r i t e r w i l l  be advanced  Maplestone,  some i n support, some c l a i m i n g v a r i o u s  The w r i t e r chooses to adhere  Anderson  and  historical  d i s c u s s why  M o d i f i c a t i o n s of t h i s  system  he will  and the r e v i s e d composition of the genera  Chandlere11a. S k r i a b i n o c t a . and S p i e n d i d o f i l a r i a w i l l  be out-  lined. 1. H i s t o r i c a l summary and s i g n i f i c a n c e of c u t i c u l a r bosses The genus Chandlere11a was  erected  (86) f o r F i l a r i a  b o s e i Chandler, 19.24; by 1949  it.. contained t h i s  o t h e r s , a l s o with non-bossate  c u t i c l e - C.  G i l b e r t , 1 9 3 0 , C. s i n e n s i s L i . 1933.  stantschinskyi  C. lepidoqrammi  and Masilungan, 1937? and. C. l i e n a l i s At  species and 4  O r i o f f , 19.47.  t h i s ' t i m e the genus S p i e n d i d o f i l a r i a  3 bossate s p e c i e s only - S. pawlowskvi  Tubangui  comprised  S k r j a b i n . 1923,  q e d o e l s t i Trav.assos., 1925.,. and S. b r e v i s p i c u l u m Singh,  S. 1949.  When he d e s c r i b e d the last-mentioned species Singh  * This species has been shown (14,59) to belong i n the Aproctinae Yorke and Maplestone, 1926.  54 (68)  commented " I t . . . appears probable that the genus  Chandlerella  i s i d e n t i c a l with S p l e n d i d o f i l a r i a " .  There i s  no way of knowing whether he was aware of a l l d e s c r i b e d and  h i s o p i n i o n was based on h i s observation  bosses  (were) d i s c e r n i b l e only i n the l i v e  difficult implied,  t o observe them i n preserved apparently,  Chandlerella  11  . . .cuticular  condition, and i t i s  material".  Thus he_  1) that some or a l l species of  a c t u a l l y possess bosses, or . 2) that the presence  of bosses should ter  either  that  species,  be.ignored_as .a generical.ly s e p a r a t i v e  since they are sometimes not v i s i b l e  same time, Singh Bucklevfilaria  (68) d e s c r i b e d  when present.  characAt the  a new genus and s p e c i e s ,  b u c k l e v i . with, "...minute, c u t i c u l a r papillae.... ' 1  which he noted, i n l i v i n g  specimens but which were " . . . d i f f i c u l t  to observe... i n f i x e d , and preserved  specimens".  ..Subsequently, Chabaud and Choquet  (24) accepted  Singh's suggested synonymization of C h a n d l e r e l l a  with  S p l e n d i d o f i l a r i a . and t r a n s f e r r e d C. lepidoqrammi to Paramicipsella  Chow, . 1 9 3 9 . I n 1957, Yen (85), on the other  s t a t e d without supporting  evidence that " C u t i c u l a r bosses  always.remain observable".  A f t e r examining Singh's  hand, should  (preserved)  type, specimens of B u c k l e v f i l a r i a b u c k l e v i . Yeh concluded that Singh had a c t u a l l y not seen "minute c u t i c u l a r p a p i l l a e " i n living  specimens, and. that  Chandlerella  "As a r e s u l t the synonymy of  and S p l e n d i d o f i l a r i a i s unacceptable" .  * Pseudaorocta b u c k l e v i  1  (Singh,  1949) Yeh, 1957  (85).  55 N e i t h e r Singh's arguments (68)  against the use  c u t i c u l a r bosses as s e p a r a t i v e c h a r a c t e r s , nor those (85)  i n favour of using them seem p a r t i c u l a r l y  of  of Yeh  l o g i c a l or very  convincing. In 1959, suppressed, and and  sub-family, the  They separated  Ornithofilariinae 10 other  Chabaud and  e x c l u s i v e groups. _Rasheed  erected a  (59)  (23). "  have attempted to show  and  C h a n d l e r e l l a formed  seems to have been the f i r s t  bosses; however, she  to l o g i c a l c o n c l u s i o n s .  She  did'not ignored  1957  not  "...constant  a t t r i b u t e s such as  absence of p r e a n a l p a p i l l a e ;. .  11  to  attached more  s p e c i e s such as Chandlere11a b r a z i l i e n s i s Yeh, conform to her generic  new  Anderson, 1959,. f o r  of the oesophagus, to which she  observations  from  G e n e r a l l y , t h e i r arguments were not  s e p a r a t i v e s i g n i f i c a n c e than'to fpllowher  and  " s h o r t - t a i l e d " genera  that the s p e c i e s of S p l e n d i d o f i l a r i a  c o n s i d e r the nature  Chandlerella  Splendidofilaria  More r e c e n t l y , most authors  very r e a l i s t i c .  left  (with 13 s p e c i e s ) *  on the b a s i s of length of t a i l  l a t t e r genus' and  mutually  (14)  (with 10 s p e c i e s ) * each genus c o n t a i n i n g  species.  Ornithofilaria  Chabaud  retained Splendidofilaria  Ornithofilaria  some bossate  Anderson and  which d i d  in • Chandlerella.  Rasheed s t a t e d "...the oesophagus of C h a n d l e r e l l a i s d i v i d e d i n t o muscular, and g l a n d u l l a r ( s i c ) portions.. Yorke and Maplestone . d e s c r i b e d the oesophagus as '.stout, short and not d i v i d e d i n t o two parts'.. Excepting these authors, a l l the others have r e p o r t e d the oesophagus as s t o u t , but not d i v i d e d i n t o two p a r t s . Hence the l a t t e r statement i s to be accepted."  * To the., combinations they l i s t e d (14, pp. 59-60) could have been added 0. c a l i f o r n i e n s i s (Wehr and Herman, 1956) and 0. bohmi Supperer, 1958.  56 This c o n c l u s i o n i s e i t h e r naive or i t was  drawn to  avoid having to r e c o n c i l e the u n d i v i d e d oesophagus of C. b o s e i (the  s p e c i e s on which  (86) t h i s genus was  based) with the  d i v i d e d oesophagus of most other s p e c i e s . In "...justified  1962,  i n r e g a r d i n g Chandlere11a  as independent  genera... ' because 1  sence or absence his  1961  Sultana (78) s t a t e d that she and  Jones  felt  Splendidofilaria  (45) had used the pre-  of c u t i c u l a r bosses as primary dichotomies i n  key to the s p e c i e s of S p l e n d i d o f i l a r i a  and. Chabaud, •.1959) .  (sensu  Anderson  It seems unnecessary to comment on t h i s  l i n e of r e a s o n i n g . N e i t h e r of the l a s t two Ornithofilaria of  d e s p i t e the s i m i l a r i t y o f . i t s  Chandlerella  and S p l e n d i d o f i l a r i a .  attached.unwarranted Ornithofilariinae Anderson  s p e c i e s to those  They seem to  have  s i g n i f i c a n c e to the removal of the  from the S p l e n d i d o f i l a r i i n a e by Chabaud and  a paper appearing a.month a f t e r Anderson's  (10), Sonin (71) used a combination of  ornamentation, relative  the genus  (23). In  revision  authors mentioned  2):oesophageal d i v i s i o n ,  and  1961  l) cuticular  3) length of t a i l  to width of anus and s p i c u l a r length, to diagnose the  three genera, i n summary, as Chandlere11a  follows:  - c u t i c l e . s m o o t h or t r a n s v e r s e l y striated' - oesophagus d i v i d e d or not - male t a i l longer than s p i c u l e s and twice as long as diameter at anus  57 Spiendidofilaria  Ornithofilaria  The  - c u t i c l e bossate - oesophagus not d i v i d e d - male t a i l l e s s than twice diameter at anus - c u t i c l e transversely striated - oesophagus d i v i d e d or not - male t a i l e q u a l l i n g diameter at anus, both exceeded by length of s p i c u l e s  p r i n c i p a l , weakness of t h i s system i s that i t  u t i l i z e s degrees of v a r i a t i o n surprising  to f i n d that  i n s t r u c t u r a l r a t i o s . I t i s not  i n t e r g r a d a t i o n s occur which tend to  negate the use of these r a t i o s  as r e p r e s e n t a t i v e  characters.  For  example, Sonin placed L o p h o r t o f i l a r i a  and  Herman, 1956 (82) i n O r n i t h o f i l a r i a . but t h i s species has  longer t a i l , than s p i c u l e s scheme a c t u a l l y  c a l i f o r n i e n s i s Wehr  (82), and according to Sonin's  should be i n c l u d e d i n C h a n d l e r e l l a  (beside  s p e c i e s with which i t c l e a r l y has much l e s s a f f i n i t y ) . well,  As  t h e . w r i t e r has specimens of an undescribed s p e c i e s of  Spiendidofilaria  (with d e l i c a t e  c u t i c l e ) , from Mexican b i r d s ,  and the males possess a t a i l more  than twice the diameter at anus. categorization,  oesophagus and bossate  On the b a s i s of Sonin's  t h i s s p e c i e s could not be placed i n  Spiendidofilaria.  f o r i t s i n c l u s i o n would i n v a l i d a t e  h i s use  of the caudal r a t i o s . In Ornithofilaria  1961, Anderson (10) suppressed the genus.. and a l l o c a t e d  its.species  which.he r e v i v e d , and S p i e n d i d o f i l a r i a .  to Chandlere11a. He.rejected the. use  of c u t i c u l a r bosses, and diagnosed these two genera on  the b a s i s of oesophageal s t r u c t u r e .  primarily  He d e s c r i b e d the  58 oesophagus  of S p i e n d i d o f i l a r i a  as " . . . l o n g , narrow,  a p p a r e n t l y devoid of g l a n d u l a r t i s s u e cated, from i n t e s t i n e " muscular, sometimes  and i m p e r f e c t l y demar-  and that of C h a n d l e r e l l a as "...broad,  clearly.divided  i n t o a n t e r i o r muscular  part and longer p o s t e r i o r , g l a n d u l a r p a r t . oesophagus  and i n t e s t i n e c l e a r l y  Junction  Anderson's concept of these genera, such as i s not w e l l enough d e f i n e d , c i e n t l y dichotomous, and  between  demarcated."  Various minor c r i t i c i s m s can be l e v e l l e d  species f a i l  undivided,  against  l ) h i s terminology  2) h i s diagnoses are not  suffi-  3) p u b l i s h e d d e s c r i p t i o n s " o f ' s e v e r a l  to mention the nature of"the"oesophagus.or i t s  j u n c t i o n , or do not seem to conform to h i s diagnoses. B. Chitwood  (personal communication,  1965)  Mrs.  M.  has drawn the '  w r i t e r ' s a t t e n t i o n to the f a c t that Anderson has been inconsistent bosses. baud  i n h i s r e j e c t i o n of the taxonomic value of c u t i c u l a r Whereas Chabaud  (14), Chabaud  and Choquet  and Anderson  (24), Anderson and Cha-  (23), and Anderson  (10)  c o n s i d e r e d the presence, or absence of c u t i c u l a r bosses to have no g e n e r i c a l l y ' s e p a r a t i v e value i n the S p l e n d i d o f i l a r i a Chandlerella-Ornithofilaria  complex, these .writers  (24, 23)  used t h i s c h a r a c t e r as the only morphological dichotomy i n d i s t i n g u i s h i n g between i n the D i r o f i l a r i i n a e example,  c e r t a i n genera of mammalian and the S p l e n d i d o f i l a r i i n a e .  the presence or absence of bosses was  to separate Loa S t i l e s .  1905 from D i r o f i l a r i a  Henry, 1910, and by Chabaud  and Anderson  filarioids For  used (24, 23) Railliet  and  (23) to separate  59 Onchocerc.ella Yorke and Maplestone,  1931  from Johnstonema  Yeh,  1957. It  i s not c l e a r  l ) whether these authors were  g u i l t y of a major o v e r s i g h t , or bosses  2) whether they thought  the  of other genera were not homologous.to those of  Splendidofilaria-Chandlere11a-Ornithofilaria separative, value, o u t s i d e t h i s complex, or  and were of  3) whether they  regarded t h i s complex as an e x c e p t i o n to general a p p l i c a b i l i t y of  presence  or absence of.bosses as.,a g e n e r i c c h a r a c t e r .  Anderson. (10, p. . 203). s t a t e d , "Chabaud and Choquet w i s e l y r e j e c t e d the use;of c u t i c u l a r bosses  (1953)  as generic  c h a r a c t e r s of f i l a r i o i d s  c l o s e to S p l e n d i d o f i l a r i a  t h o f i l a r i a ..."  added;by the present w r i t e r ) .  (emphasis  and O r n i This  o p i n i o n by Anderson might be construed as i n d i c a t i n g that use of bosses the  should not n e c e s s a r i l y be r e j e c t e d  the  throughout  Filarioidea, The w r i t e r has not examined bossate mammalian  filarioids  and i s t h e r e f o r e not i n a p o s i t i o n to express  an  o p i n i o n on the s i g n i f i c a n c e ; of these s t r u c t u r e s i n that  group.  N e v e r t h e l e s s , the w r i t e r agrees with Anderson (10) that  the  presence  of the narrow, s i m p l i f i e d  Splendidofilaria-Ornithofilaria more important  than, the presence  type of oesophagus i n the  complex should be.considered or absence of bosses,  and  that there are. too. many s i m i l a r i t i e s between t h e / s p e c i e s of Splendidofilaria  and O r n i t h o f i l a r i a with t h i s type, of oesopha-  gus t o permit r e c o g n i t i o n of O r n i t h o f i l a r i a genus.  as an  independent  The the  w r i t e r does not b e l i e v e that the presence of  usual types of f i l a r i o i d  similarities fies  oesophagus i n other  l i e i n t h e i r lack of d i s t i n c t i v e  features,  Anderson's i m p l i c a t i o n (10) that C h a n d l e r e l l a  e q u a l l y homogeneous, and e q u a l l y r e a l i s t i c the  contrary,  vising  generic  species.  genus S k r i a b i n o c t a  of i n f o r m a t i o n The w r i t e r w i l l  provided  i s an On  needs r e -  by r e c e n t l y  show, why he t h i n k s the  Chertkova, 1946 must be removed from.  Chandlere11a and, haying made, t h i s change, w i l l nature, of t h i s genus by arranging  n a t u r a l and a r t i f i c i a l characters.  justi  group.  the w r i t e r t h i n k s that C h a n d l e r e l l a  i n the. l i g h t  described  the  species whos  demonstrate  i t s many species i n  groups on the. b a s i s of combinations of  I t w i l l be f u r t h e r shown that C. p e r i a r t e r i a l i s  ( C a b a l l e r o , .1948) Rasheed, I960- should Parachandlerella  C a b a l l e r o , . .1948  be returned  to .  and that C. s h a l d v b i n i  Gubanov, 1954 does not belong i n the Onchpcercidae. ' Subsequently the genus S p i e n d i d o f i l a r i a w i l l be brought up to date, and the. w r i t e r w i l l use. .his on  s e v e r a l species  diagnosis 2.  observations  to i n d i c a t e that minor changes i n i t s  are warranted.  The genus S k r i a b i n o c t a  Chertkova, 1946  This genus was e r e c t e d  on the b a s i s of specimens  from the. eye of. a dove i n the U.S.S.R. (28). Chertkova. was i n f l u e n c e d by the s i t e s t a t e d that S k r i abinocta  Evidently  of these worms when she  "...the genus most c l o s e l y  approaching  nov. gen. i s the genus Aprocta Linstow, 1883".  61 A f t e r comparing Chertkoya's d e s c r i p t i o n of Sk. specimens which he  named C h a n d l e r e l l a  ( l l ) proposed t h a t . " s i n c e r e l i a b l e and  s i d e r e d a synonym of the  latter..."  Two 1961  and  Chandlerella  found  former i s con-  Chandlere11a s t r i a t o s p i c u l a H i b l e r , 1964, (Chertkova, 1946)  chitwoodae Anderson, 1961.  closely  Anderson, 1961  Tables I and  II (pp.  Jones,  and  C.  21-24) i n d i -  only minor d i f f e r e n c e s i n q u a n t i t a t i v e f e a t u r e s among  these 4 s p e c i e s .  As w e l l , they are very  following q u a l i t a t i v e characters: cuticle,  anus and  caudal  p a p i l l a e of male, and  extremity  type of  of t h i s group of 4 species Skriabinocta  should  oesophagus of the striatospicula  of female, s p i c u l e s ,  i s not  characters  s i m i l a r , and  2) i t has  the w r i t e r b e l i e v e s that the  The  The  genus  divided  chitwoodae. f l e x i v a g i n a l i s .  as undivided  of these species  and  are  (28)  because  strikingly  been shown (9) that f o r at l e a s t  ( D i p l o t r i a e n a spp.)  used f o r generic  nature  a s e r i o u s b a r r i e r to p l a c i n g them with  other  some f i l a r i o i d s  distinctive  be r e v i v e d to c o n t a i n them.  species  caudal  microfilaria.  petrowi i n which i t i s d e s c r i b e d  l) the  s i m i l a r i n the  cephalic p a p i l l a e ,  In view of the homogeneous and  Sk.  the  cannot be  species, S p l e n d i d o f i l a r i a f l e x i v a g i n a l i s  resemble C. petrowi  cate  chitwoodae. Anderson  characters  t o separate S k r i a b i n o c t a  petrowi with  t h i s character  cannot  be  separation.  available description  l i e n a l i s O r l o f f , 1947  embodies the  of Chandlere11a  f o l l o w i n g f e a t u r e s which  62 suggest a f f i n i t y with S k r i a b i n o c t a :  4 p a i r s of  p a p i l l a e , sabrelike s p i c u l e s , abruptly tail,  and  no  female anus.  tentatively,  in  The 1946  The  c o n s t r i c t e d female  w r i t e r places  C.  lienalis.  Skriabinocta. diagnosis  of the genus S k r i a b i n o c t a  i s amended to read as f o l l o w s :  C u t i c l e smooth and  thin.  Chertkova,  Splendidofilariinae.  Extremities  rounded.  c e p h a l i c p a p i l l a e , r e g u l a r l y arranged i n two amphids.  cephalic  Four p a i r s  circles,  Oesophagus moderately broad, d i v i d e d or  plus  undivided.  Anus of female s c a r c e l y d i s c e r n i b l e , u s u a l l y d i s p l a c e d T a i l of female u s u a l l y abruptly  narrowed and  slightly  expanded, rounded knob.  slightly  d i s s i m i l a r , pointed,  strongly  weakly developed capitulum and of postanal structure  p a p i l l a e ; and  on  Type Sk.  terminating  simple,  arcuate,  calomus.  in  subequal,  and  with  Three to seven p a i r s  u s u a l l y , a simple  a n t e r i o r l i p of c l o a c a .  c y s t s beside blood v e s s e l s of b i r d s .  Spicules  dextrad.  papilla-like  Coiled in  thin-walled  of body c a v i t y or w i t h i n  the  eyeball,  M i c r o f i l a r i a e b l u n t - t a i l e d , i n blood stream.  species: petrowi Chertkova, 1946 Syn.  S p l e n d i d o f i l a r i a petrowi Anderson and -Chandlerella  Chabaud, 1959 petrowi  Anderson, 1961 Host:  (28) (Chertkova, (14)  (Chertkova,  1946)  (ll)  S t r e p t o p e l i a o r i e n t a l i s meena (Sykes) (Columbidae)  1946)  63 Site:  I n t e r n a l media of eye adherent t o v a s c u l a r membrane  Locality: Other  Kirghizia,  U.S.S.R.  species:  Sk. l i e n a l i s Syn.  (Orloff,  1947) n. comb.  Chandlerella  lienalis Orloff.  Paramicipsella  lienalis  Chabaud and Choquet, Ornithofilaria Anderson,  1947  (69)*  ( O r l o f f , 1947)  1953 (24)  lienalis  ( O r l o f f , 1947)  1955 (4)  Parachandlere11a l i e n a l i s  ( O r l o f f , 1947)  S u l t a n a , 1962 (78) Host:  Alectoris  Site:  Splenic  Locality:  Armenia  Sk. chitwoodae Syn.  qraeca (Meisner) (Phasianidae)  vessels  (Anderson, 1961) n. comb.  Chandlere11a  chitwoodae  Anderson,  1961  Host:  Padda o r v z i v o r a  Site:  In lungs, probably i n blood v e s s e l s  Locality:  O n t a r i o , Canada, imported from Java  Sk. f l e x i v a q i n a l i s Syn.  (ll)  (L.) (Ploceidae)  (Jones, 1961) n. comb.  Spiendidofilaria Chandlere11a  flexivaqinalis  flexivaqinalis  Jones, 1961 (45)  (Jones, 1961)  S u l t a n a , 1962  * P u b l i s h e d i n F i l a r i a e of Animals and Man (69) without r e f e r e n c e to a paper by O r l o f f .  64  Hosts  (Corvidae): 1)  Hosts  Site:  Corvus b. brachyrhynchos Brehm (45)  (Tetraonidae): 2)  Dendragapus obscurus f u l i g i n o s u s  3)  Dendragapus obscurus s i t k e n s i s  4)  Dendragapus obscurus p a l l i d u s  5)  Dendragapus obscurus r i c h a r d s o n i  6)  Bonasa umbellus  7)  Bonasa umbellus brunnescens  8)  Canachites canadensis f r a n k l i n i  (Ridgway)  Swarth Swarth (Douglas)  a f f i n i s A. & F. Conover (Douglas)  In t h i n - w a l l e d mesenteric or p e r i t o n e a l throughout the body c a v i t y , u s u a l l y with a major  cysts  associated  a r t e r y ; or w i t h i n the adrenal  glands. Localities:  l) 2-8)  Sk. s t r i a t o s p i c u l a Syn.  Ohio, U.S.A. (45), and B r i t i s h Columbia,  ( H i b l e r , 1964) n. comb.  C h a n d l e r e l l a s t r i a t o s p i c u l a H i b l e r , 1964 (44)  Host:  P i c a p i c a hudsonia  Site:  Connective t i s s u e around s p l e n i c oesophagus,  Locality: 3.  Canada  (Sabine) (Corvidae) artery,  or i n the mesenteries.  Colorado, U.S.A.  The genus C h a n d l e r e l l a With the removal of S k r i abinocta from t h i s  the species p l a c e d i n i t by Anderson recently f a l l  i n t o 4 groups.  genus,  (10) and those d e s c r i b e d  A l l these species are separable  65 from those  of S p l e n d i d o f i l a r i a  bv t h e i r robust  The c h a r a c t e r s of a l l groups are presented  oesophagus.  first,  then f o r  each group, i n t u r n , the s p e c i e s on which the groupings based w i l l be l i s t e d  are  and d i s c u s s e d .  Group A Divided oesophagus; r a t h e r s h o r t , s t o u t , b l u n t - t i p p e d s p i c u l e s ; long d i g i t i f o r m male t a i l (more than twice as long as the s p i c u l e s ) ; no preanal p a p i l l a e , and 3-5 p a i r s of p o s t a n a l s . Group B Undivided oesophagus; curved pointed s p i c u l e s ; modera t e l y long male t a i l ; no p r e a n a l p a p i l l a e , and s e v e r a l p a i r s of p o s t a n a l s . Group C D i v i d e d oesophagus; s p i c u l e s curved, u s u a l l y p o i n t e d ; short male t a i l (about same length as s p i c u l e s ) ; p r e a n a l p a p i l l a e sometimes present, and s e v e r a l p a i r s of p o s t a n a l s . Group D Undivided oesophagus; curved, pointed s p i c u l e s ; short male t a i l ; few caudal p a p i l l a e ; c u t i c l e bearing bosses. Species  of Group A  C. c o l u m b i q a l l i n a e (Augustine, 1937) Rasheed, 1960 (59). Syn. V a g r i f i l a r i a c o l u m b i q a l l i n a e (15), S p l e n d i d o f i l a r i a c o l u m b i q a l l i n a e (Augustine. 1937) Chabaud and Choquet, 1953 (24). Hosts: C o l u m b i g a l l i n a p a s s e r i n a n i g r i r o s t r i s (15), (?) Crocopus phaenicooterus (59)(Columbidae). S i t e : Body c a v i t y (15, 5 9 ) , h e a r t , l i v e r , pulmonary v e s s e l s (15). L o c a l i t i e s : V i r g i n Islands (15), (?) India (59), C. s i n e n s i s L i , 1933 (47). Syn. S p l e n d i d o f i l a r i a s i n e n s i s T L i , 1933) Chabaud and Choquet, 1953 (24), Hosts: U r o c i s s a s i n e n s i s . Corvus s i n e n s i s (47), Corvus macrorhvnchus (2) ( C o r v i d a e ) ; ( ?)Di'crurus f orf i c a t u s (26) (Dicruridae) . S i t e : Lung, 'tracheal (47) , l i v e r (26).. L o c a l i t i e s : China (47), India (2), (?)Malgache Republic (26).  66 C. s i n o h i A l i , 1956 (2). Syn. S p i e n d i d o f i l a r i a sincrhi .(Ali, 1956) Anderson and Chabaud, 1959 (14). Host: Caprimulous sp. (C. a s i a t i c u s according to Rasheed (59)) (Caprimulgidae). S i t e : Heart. L o c a l i t y : India. C.  t h a p a r i Rasheed, 1960 (59). Host: S a x i c o l a torouata S i t e : Body c a v i t y . L o c a l i t y : India.  indica  (Turdidae).  C. himalavensis S u l t a n a , 1962 (78). Host: Cerchneis tinnunculus i n t e r s t i n c t u s - S i t e : Heart and p e r i c a r d i a l c a v i t y . . L o c a l i t y : India. C.  Sultana, 1962 (78). Host: M o t a c i l l a maderaspatensis S i t e : Heart. L o c a l i t y : India.  (Falconidae).  alii  (motacillidae).  C. suitanae nomen novum f o r C. s t a n t c h i n s k v i - G i l b e r t , 1930 of Sultana 1962 (78~T Host: Uroloncha s t r i a t a s t r i a t a (Ploeeidae). S i t e : Heart. L o c a l i t y : India (78). Sultana's "C. c e r t a i n l y not  the  appears to be  a distinct  others short  s t a n t c h i n s k v i G i l b e r t . 1930"  species which G i l b e r t (36)  described.  s p e c i e s , c l e a r l y separable  of t h i s group by i t s narrow diameter, short oesophagus, and  other  is It  from  the  female  measurements i n combination  tail,  (78);  t h e r e f o r e , the w r i t e r proposes f o r i t the p r o v i s i o n a l r e p l a c e ment name C h a n d l e r e l l a Group A may Vaqrifilaria leave  suitanae  nomen novum.  deserve generic  Augustine, 1937;  status,  as  however, the w r i t e r p r e f e r s  i t i n Chandlere11a at present.  Group A seems to be a  n a t u r a l assemblage of c l o s e l y r e l a t e d s p e c i e s , and forms a homogeneous nucleus around which can be s p e c i e s which lack w e l l d e f i n e d  to  separative  as  such,  assembled  characters.  It  the  67 could be argued t h a t the homogeneity which appears to e x i s t among the 7 s p e c i e s of Group A a c t u a l l y i n d i c a t e s a smaller number of r a t h e r v a r i a b l e s p e c i e s , some of the geographic or host-dependent v a r i a t i o n s of which have been given status.  Indeed, C. s i n q h i A l i , 1956 i s very  s i n e n s i s L i , 1933. of specimens  specific  s i m i l a r t o C.  I t i s obvious that c r i t i c a l  re-examination  ( i n c l u d i n g m i c r o f i l a r i a e ) and a c q u i s i t i o n of more  abundant m a t e r i a l are necessary before  w e l l founded  conclu-  s i o n s can be drawn. The be placed  remaining 12 species which have been, or could  i n Chandlerella  which i s separable  form a very heterogeneous assemblage  from other  sence of c h a r a c t e r s .  genera p r i n c i p a l l y by the ab-  These species  f e a t u r e s and many d i f f e r e n c e s .  show few common p o s i t i v e  Two s p e c i e s , C. o e r i a r t e r i a l i s  and  C. s h a l d v b i n i . which have been placed  38)  should  be removed, and S p l e n d i d o f i l a r i a g r e t i l l a t i  baud, Anderson, and Brygoo, 1959 should Species  i n t h i s genus (59, Cha-  be t r a n s f e r r e d i n t o i t .  of Group B  C. b o s e i (Chandler, 1924) Yorke and Maplestone, 1926 (86). Syn. F i l a r i a b o s e i (27), S p l e n d i d o f i l a r i a bosei (Chandler, 1924) Chabaud and Choquet, 1953 (24). Hosts: Dicrurus paradiseus ( D i c r u r i d a e ) (27); Falco t i n u n c u l u s ( s i c ) (Falconidae) (46); Certhia f a m i l i a r i s ( C e r t h i i d a e ) (46); Corvus f r u q i l e g u s (Corvidae) (46), "crows" (57) . S i t e : Body c a v i t y (27), a i r sac or body c a v i t y (46), heart (57). L o c a l i t i e s : India (27, 57), U.S.S.R. (46). C. g u i s c a l i (Linstow, 1904) n. comb. Syn. F i l a r i a g u i s c a l i (49). S p l e n d i d o f i l a r i a g u i s c a l i (Linstow, 1904) Odetoyinbo and Ulmer, 1960 (53).  Host: Quiscalus a u i s c u l a S i t e : V e n t r i c l e s of b r a i n L o c a l i t y : U.S.A. (49, 5 3 ) .  ( i c t e r i d a e ) (49, 53)* (49, 53).  C h a n d l e r e l l a q r e t i l l a t i (Chabaud, Anderson and Brygoo, 1959) Sonin, 1961 (71) . Syn. S p i e n d i d o f i l a r i a q r e t i l l a t i (25.) . Host: Aqapornis cana ( P s i t t a c i d a e ) (25) . S i t e : ?Body c a v i t y . L o c a l i t y : Madagascar. Chandlere11a apusi Sonin, 1963 (74). Host: Hirundapus caudacutus .(Micropidae) . S i t e : Heart and large v e s s e l s . L o c a l i t y : Amur,'U.S.S.R. Chandler  (27) s t a t e d that the female which he  d o u b t f u l l y placed with h i s male F i l a r i a phic.  Pandit  bosei was amphidel-  e_t a_l. (57) b r i e f l y r e d e s c r i b e d  C. bosei  from  a l a r g e r number of specimens and concluded that Chandler's > female d i d not belong to t h i s  s p e c i e s ; however, they  failed  to mention whether t h e i r females were amphidelphic or opisthodelphic. Chandlere11a i t s uterine  Since  C. b o s e i i s the type species of  (86) an attempt should  be made to determine  arrangement, f o r a l l the other  species which seem  to belong i n t h i s genus are o p i s t h o d e l p h i c . The described;  c e p h a l i c p a p i l l a e of C. b o s e i have not been  however, 10 of the 12 species  f o r which c e p h a l i c  p a p i l l a e have been mentioned possess 2 p a i r s , and i t seems l i k e l y that t h i s number i s t y p i c a l of the genus. t i o n s are C h a n d l e r e l l a  s h a l d y b i n i Gubanov, 1954* from the  kidney of Phalacrocorax u r i l e apusi.  According  The excep-  to Sonin  (Phalacrocoracidae),  and C.  (74), C. s h a l d v b i n i has 6 small  * Named, but not d e s c r i b e d , i n 1954 (38). Sonin's (74) seems to be the f i r s t p u b l i s h e d d e s c r i p t i o n .  69 and  4 l a r g e r c e p h a l i c p a p i l l a e , 3 p a i r s of caudal  all  preanal,  and  embryonated eggs.  Obviously,  papillae -  this  species  must be removed from the Onchocercidae; probably i t belongs i n the Aproctinae is  Yorke and  Maplestone, 1926.  C.  apusi i s  s a i d to have 6 p a i r s of c e p h a l i c p a p i l l a e (74).  this  species  c l o s e l y resembles C. b o s e i i t should  termined whether the i n the  type s p e c i e s  c o n f i g u r a t i o n of the  Since be  de-  cephalic papillae  as w e l l , i s at variance  with usual  pat-  tern. S p l e n d i d o f i l a r i a g r e t i l l a t i possesses a l l the characters  of the w r i t e r ' s Group B, and  resemblance to the Splendidofilaria Sonin (25) but  (71)  other  (10).  species  bears  i n c l u d e d by Anderson i n  Therefore,  the w r i t e r agrees with  that i t belongs i n C h a n d l e r e l l a .  d i d not mention the their illustration  little  Chabaud e_t a l .  c e p h a l i c p a p i l l a e of t h i s  species,  s t r o n g l y suggests the presence of 4  pairs. The guiscali  w r i t e r has  (Linstow, 1904)  n. comb., and  small c e p h a l i c p a p i l l a e . with 4  The  C.  found 4 p a i r s of  w r i t e r b e l i e v e s that  (or more) p a i r s of c e p h a l i c p a p i l l a e should  included  i n the  same genus with species possessing  pairs.  However, a new  present  time f o r C.  because  examined specimens of  l ) the  Chandlerella  genus w i l l not  g u i s c a l i . C.  not  a p u s i . and  2) Dr. M.  C.  be  only  be proposed at  c e p h a l i c p a p i l l a e of the  are unknown, and  species  2  the  oretillati  type species J . Ulmer, who  of sent  70 the  w r i t e r specimens of C.  his  intention  a u i s c a l i . has  of p u b l i s h i n g f u r t h e r on  indicated  this  [in  litt.)  species.  Species of Group C C. s t a n t s c h i n s k v i G i l b e r t , 1930 (36). Syn. Paramicipse11a stantchinskv ( G i l b e r t , 1932) Chabaud and Choquet, 1953 (24), O r n i t h o f i l a r i a s t a n t s c h i n s k v i ( G i l b e r t , 1930) Anderson, 1955 (4). Hosts: O r i o l u s o r i o l u s ( O r i o l i d a e ) (36); Corvus corax (Corvidae) (69). S i t e : Body c a v i t y . L o c a l i t y : U.S.S.R. (36, 69). C. p e r i a r t e r i a l i s ( C a b a l l e r o , 1948) Rasheed, 1960 (59). Syn. Parachandlere11a p e r i a r t e r i a l i s (21), Spiendidofilaria p e r i a r t e r i a l i s (CabaTlero. 1948) Chabaud and Choquet, 1953 (24) . Host: Tyrannus sp. (Tyrannidae) (21). S i t e : Connective t i s s u e over pulmonary a r t e r i e s . L o c a l i t y : Mexico. C. b r a z i l i e n s i s Yeh, 1957 (85). Syn. Spiendidofilaria b r a z i l i e n s i s (Yeh. 1957) Anderson and Chabaud, 1959 (14), P a r a c h a n d l e r e l l a b r a z i l i e n s i s (Yeh, 1957) S u l t a n a , 1962 (78). Host: Rhamphastus d i c o l o r u s (Rhamphastidae) (85). S i t e : C e r v i c a l a i r sac. L o c a l i t y : Brazdl. C.  b u c k l e v i S u l t a n a , 1962 (78). Host: L e i o o i c u s mahrattensis b l a n f o r d i S i t e : Heart c a v i t y . L o c a l i t y : India. C.  p e r i a r t e r i a l i s has  a pair  (Picidae).  of.preanal  papillae  a s . w e l l as t h e . p o s t a n a l s , t e r m i n a l caudal protuberances, "lightly chitinized" amphidelphic most of the 4 pairs and  (21).  spicules,  s p e c i e s of C h a n d l e r e l l a .  a striated  c u t i c l e , and  In a l l these c h a r a c t e r s i t d i f f e r s  of p r e a n a l p a p i l l a e , cuticle  t i e s Sultana  (78)  Orloff,  i n the  1947  a striated  (85).  C.  braziliensis  "lightly chitinized" On  the  b a s i s of these  p l a c e d these 2 species and  C.  is from has  spicules, similari-  lienalis  r e v i v e d genus P a r a c h a n d l e r e l l a  Caballero,  71 1948.  The present w r i t e r b e l i e v e s that the combination  Parachandlere 11a p e r i a r t e r i a l i s C a b a l l e r o , 1948 i s v a l i d , but that C. b r a z i l i e n s i s  should not be placed with P.  p e r i a r t e r i a l i s because of the l a t t e r ' s amphidelphism and p a i r e d caudal protuberances. diversity  In view of the morphological  e x h i b i t e d by the other s p e c i e s of C h a n d l e r e l l a .  C. b r a z i l i e n s i s  i s r e t a i n e d i n t h i s genus.  C. b u c k l e v i . with rounded s p i c u l e s , and C. s t a n t s c h i n s k v i may be r a t h e r c l o s e l y  related  of Group A, d i f f e r i n g from them p r i n c i p a l l y  to the species i n t h e i r short  tails. Species of Group D C.  s k r i a b i n i (Petrov and Chertkova, 1947^ Anderson, 1961 Svn. C a r d i o f i l a r i a s k r i a b i n i (58) , O r n i t h o f i l a r i a s k r i a b i n i (Petrov and Chertkova, 1947) Anderson and Chabaud, 1959 (14). Host: H y p o t r i o r c h i s subbuteo (Falconidae) (58), S i t e : Heart. L o c a l i t y : K i r g h i z i a , U.S.S.R.  Tl07"]  C. t r a v a s s o s i (Koroliowa, 1926) Anderson, 1961 (10). Syn. S p l e n d i d o f i l a r i a t r a n v a s s o s i (46), P a r a m i c i p s e l l a t r a v a s s o s i (Koroliowa. 1926) Chabaud and Choquet, 1953 (24), Ornittiof i l a r i a t r a v a s s o s i (Koroliowa, 1926) Anderson, 1955 ( 4 ) . Hosts: Meroos a p i a s t e r (46, 72), Merops s u p e r c i l i o s u s (72)(Coraciidae). S i t e : Heart (46, 72) and aorta (72). L o c a l i t y : U.S.S.R. L  C.  skri abini  l a c k s caudal p a p i l l a e but possesses  caudal bosses and a short b u c c a l capsule.  I t s presence i n  Chandlere11a i n c r e a s e s the h e t e r o g e n e i t y of t h i s genus, but at present  i t can not be p l a c e d elsewhere s a t i s f a c t o r i l y .  * According to Petrov and Chertkova 1950 (58) t h i s s p e c i e s was f i r s t d e s c r i b e d i n 1948 i n F i l a r i a e of Animals and Man (69).  72  C.  t r a v a s s o s i has  of preanal  a g e n e r a l l y bossate c u t i c l e ,  p a p i l l a e and  2 p a i r s of p o s t a n a l s .  and  one  Were i t not  f o r i t s broad oesophagus t h i s species might be placed Splendidofilaria;  since the  to be of b a s i c s e p a r a t i v e C h a n d l e r e l l a . there from the  nature of the  value  i s no way  l a t t e r genus at t h i s  and  C.  follows:  C.  braziliensis.  these groups was has  is  an a r t i f i c i a l  C.  columbiqallinae.  He  C_.  s i n e n s i s . C.  so, but  lienalis.  The  writer  that groupings can  be  (the w r i t e r ' s Group  I t i s apparent that Chandlere11a  assemblage of s e v e r a l genera.  u n t i l new  into  implied that each of  natural a f f i n i t i e s  significant: characters  the  sinqhi.  p o s s i b l y , Group B).  mined, and  genus C h a n d l e r e l l a  "...remarkably homogeneous..."  made which r e f l e c t and  the  s t a n t s c h i n s k v i . C.  and  travassosi  Group A - C. b o s e i . C. t r a v a s s o s i .  shown that t h i s i s not  A,  i n S p l e n d i d o f i l a r i a and  time.  skr i a b i n i : Group B - C.  p e r i a r t e r i a l i s . C.  in  oesophagus seems  of removing C.  Anderson subdivided 2 groups, as  pair  of the type,  Until  C. b o s e i . are  d e s c r i p t i o n s can be used to  deterclarify  i n t e r r e l a t i o n s h i p s of i t s s p e c i e s , Chandlere11a  be r e t a i n e d i n i t s heterogeneous s t a t e .  The  the  should  w r i t e r regards  Chandlere11a as a convenient, temporary r e p o s i t o r y f o r avian  s p l e n d i d o f i l a r i i n e nematodes with a broad oesophagus  of moderate length,  a preoesophageal v u l v a ,  no  distinctive  c e p h a l i c s t r u c t u r e s , subequal, s l i g h t l y d i s s i m i l a r s p i c u l e s ;  73 u s u a l l y with no buccal capsule or bosses, and with caudal p a p i l l a e i n l o n g i t u d i n a l rows; and o f t e n with only 2 p a i r s of  cephalic 4.  papillae.  The genus S p l e n d i d o f i l a r i a The  Anderson's  f o l l o w i n g s p e c i e s have been d e s c r i b e d  since  1961 r e v i s i o n :  ( C u t i c u l a r bosses present) S. s i n a h i S u l t a n a , 1962. Host; - M o t a c i l l a maderaspatensis S i t e : Subcutaneous. L o c a l i t y : India (78). S. a l i i  S u l t a n a , 1962. Host: Leptocoma a s i a t i c a S i t e : Heart. L o c a l i t y : India (78).  (Motacillidae) .  asiatica  (Nectariniidae).  S. p i c a c a r d i n a H i b l e r , 1964. Host: Pic a p i c a hudsonia (Corvidae). S i t e : Heart. L o c a l i t y : Colorado, U.S.A. (44). ( C u t i c u l a r bosses  absent)  S. caperata H i b l e r . 1964 Host: Pic a p i c a hudsonia (Corvidae). S i t e : Heart. L o c a l i t y : Colorado, U.S.A. (44). S. p e c t o r a l i s n. sp. Hosts: Dendraoapus obscurus. Bonasa umbellus. Canachites canadensis. Pedioecetes phasianellus(Tetraonidae). S i t e : Subcutaneous. L o c a l i t i e s : B r i t i s h Columbia, Canada; A l a s k a , U.S.A. ( S p l e n d i d o f i l a r i a sp. A. Host: Dendraoapus obscurus (Tetraonidae). S i t e : Connective t i s s u e near oesophagus. L o c a l i t y : B r i t i s h Columbia, Canada.) The w r i t e r attempted to determine whether the s p e c i e s of S p l e n d i d o f i l a r i a  sensu Anderson  1961 (10) could  74 be segregated  i n t o n a t u r a l groups by using v a r i o u s  t i o n s of c h a r a c t e r s . s p e c i e s and  combina-  In a d d i t i o n to specimens of the  S. p a p i l l o c e r c a from l o c a l  new  t e t r a o n i d s and  pub-  l i s h e d d e s c r i p t i o n s of the other s p e c i e s , the w r i t e r examined specimens of S. S.  fallisensis  aloonquinensis  undescribed  (Anderson, 1954)  (Anderson, 1955)  Anderson,  Anderson, 1961,  species s i m i l a r to S. wehri Anderson,  A f t e r comparing c h a r a c t e r s between bossate (Spiendidofilaria (Ornithofilaria  sensu Sonin  sensu S o n i n ) ,  groups, the w r i t e r reached the l ) No  evident  (71))  and  an  1961.  species  non-bossate  and w i t h i n each of  species  these  following conclusions:  c o r r e l a t i o n e x i s t s among any com-  b i n a t i o n s of the presence, absence, or degree of of the  and  1961,  following characters:  bosses or other  expression  conspicuous  c u t i c u l a r elements, p a i r e d caudal protuberances,  precloacal  protuberance, caudal p a p i l l a e , s p i c u l a r s t r u c t u r e , (or s i t e , f a m i l y of host,  or l o c a l i t y ) .  s p e c i e s have been recovered  However, most of the  bossate  from the heart, whereas most of  the non-bossate species have been found subcutaneously, .2-) The s p e c i e s , ranging (3) and  S.  s p i c u l e s are somewhat v a r i a b l e between from short and d e l i c a t e i n S.  aloonquinensis  i n S. p e c t o r a l i s n. sp. which the  fallisensis  (4) to moderately long and However, f o r the  13 species i n  s t r u c t u r e of the s p i c u l a r apices and the  s p i c u l a r lengths  robust  can be determined, the t i p of the  relative left  s p i c u l e i s more s h a r p l y p o i n t e d than that of the r i g h t ,  and  75 the is  left  s p i c u l e i s longer than the r i g h t .  S. t u v e n s i s  (76), i n which the r i g h t  e n t l y more s h a r p l y p o i n t e d , and 3)  The  (The  exception  s p i c u l e i s appar-  i s longer than  c u t i c u l a r r i d g e s and unusual  the  left.)  cephalic  compression i n S. p a p i l l o c e r c a and S p i e n d i d o f i l a r i a  sp. A  are of s e p a r a t i v e value only at the s p e c i e s l e v e l s i n c e a) S. caperata H i b l e r , 1964 c u t i c l e but  (44) possesses  t h i s type  lacks the c e p h a l i c compression, and  papillocerca  b)  i s very s i m i l a r i n other r e s p e c t s to  p e c t o r a l i s which lacks both of these  characters.  of S.  S. It i s  u n l i k e l y that the c e p h a l i c compression i s a f i x a t i o n a r t e f a c t because the Alaskan  specimens of S. p a p i l l o c e r c a were  f i x e d i n a l c o h o l - f o r m a l i n - a c e t i c f i x a t i v e , the S o v i e t specimens i n "Barbagallo's  solution"  (M. D. Sonin, i n  l i f t . ) . and the male S p i e n d i d o f i l a r i a 4) Contrary a e d o e l s t i Travassos, (Oschmarin, 1950) rotundicephala  ethanol.  to Anderson's o p i n i o n (10),  1925  (79)  and S.  Anderson, 1961,  (56), probably  S. o e d o e l s t i was  sp. A i n 70%  rotundicephala  syn.  Ornithofilaria  do not belong  d e s c r i b e d as having  S,  i n t h i s genus.  a small v e s t i b u l e , and  an oesophagus d i f f e r e n t i a t e d  i n t o a pharynx i n i t s a n t e r i o r  third;  a d i g i t i f o r m male t a i l  as w e l l , i t possesses  None of these of  this  c h a r a c t e r s i s shared with any  large group.  other  Because of i t s apparently  (79).  species  narrow  oesophagus the w r i t e r p r e f e r s to leave i t p r o v i s i o n a l l y i n Spiendidofilaria  sensu l a t o . r a t h e r than to t r a n s f e r i t to  76 a new  taxon or to C h a n d l e r e l l a . The  unusually  broad and uniform  p l a c e d with The  oesophagus of S. r o t u n d i c e p h a l a  seems of  diameter to allow i t to be  such species as S.  fallisensis  and S.  pectoralis.  extremely long oesophagus of S. r o t u n d i c e p h a l a  site  ( w i t h i n the o r b i t ) may  ties  l i e elsewhere, but  Splendidofilaria 5)  a l s o i n d i c a t e that i t s a f f i n i -  the w r i t e r leaves i t i n  sensu l a t o f o r the time  The  and i t s  oesophagi of those  being.  species which have  been d e s c r i b e d or f i g u r e d i n some d e t a i l seem to be similar.  very  In the 6 s p e c i e s examined by the w r i t e r the  phagus i s moderately long, very narrow, d e l i c a t e , and developed.  The  a n t e r i o r narrowest r e g i o n has  oesoweakly  narrow  homogeneously h y a l i n e w a l l s apparently without  nuclei,  and  a very narrow lumen the c u t i c u l a r l i n i n g of which i s sometimes sinuous.  The  appears v a c u o l a t e , pass forward  and  immediately p o s t o r a l p o r t i o n f r e q u e n t l y and  around i t strands of  outward to the  (••?) n e u r a l t i s s u e  surface of the head.  p o s t e r i o r component of the oesophagus i s s l i g h t l y and more g r a n u l a r than the a n t e r i o r , and broader lumen.  broader  a slightly  This p o r t i o n i s somewhat v a r i a b l e between  i n d i v i d u a l s of any  one  s p e c i e s , and  t i o n i n S. p a p i l l o c e r c a and to be  has  The  shows g r e a t e s t  elabora-  S. p e c t o r a l i s where i t appears  a very rudimentary g l a n d u l a r oesophagus.  t u d i n a l o p t i c a l s e c t i o n the w a l l s of t h i s r e g i o n  In  longi-  comprise  an outer non-nucleate l a y e r of i r r e g u l a r o u t l i n e , and  an  77 i n n e r l a y e r which i s broader, m u l t i n u c l e a t e , lar.  more granu-  A short t r a n s i t i o n a l zone u s u a l l y i s evident  the  two  the  i n t e s t i n e , although the  oesophageal regions  and  between the  these two was  The  h i s t o l o g i c a l l y d i f f e r e n t regions  noted i n a l l 6 s p e c i e s  illustrations picacardina  (44)  and  S.  between  oesophagus  i n t e s t i n e i s sometimes  i n f l a t e d where the oesophagus j o i n s i t .  gus  and  abruptly  existence  of the  examined, and  and  of  oesopha-  Hibler's  suggest a s i m i l a r s i t u a t i o n i n S. caperata.  In c o n c l u s i o n ,  i t appears that with the  removal  of S. q r e t i l l a t i  from t h i s genus to Chandlere11a and  r e l e g a t i o n of S.  q e d o e l s t i and  p o s i t i o n outside  the major group, the remaining  S. r o t u n d i c e p h a l a  form a reasonably homogeneous assemblage on the  the  to a species basis  of  oesophageal s t r u c t u r e . Anderson's amended d i a g n o s i s Spiendidofilaria  (10)  of the  genus  reads:  "Splendidofilariinae. Oesophagus long, narrow, undivided, apparently devoid of g l a n d u l a r t i s s u e and i m p e r f e c t l y demarcated from i n t e s t i n e . S p i c u l e s subequal but d i f f e r e n t i n morphology. C u t i c l e sometimes with bosses. Vulva ant e r i o r to end of oesophagus. Caudal p a p i l l a e o f t e n much reduced, u s u a l l y i n rows on v e n t r o l a t e r a l surface of caudal extremity. Two p a i r s of c e p h a l i c p a p i l l a e and amphids present, o f t e n arranged a s y m e t r i c a l l y about stoma. Tail u s u a l l y t e r m i n a t i n g i n a p a i r of f l e s h y protuberances (not phasmids). M i c r o f i l a r i a s h o r t . P a r a s i t e s of the blood v a s c u l a r system, subcutaneous t i s s u e s , and serous c a v i t i e s of b i r d s . " The. w r i t e r p r e f e r s to :amend. Anderson's as  follows:  Splendidofilariinae.  long, narrow, and  diagnosis  Oesophagus moderately  d e l i c a t e ; i t s very  narrow, h y a l i n e ,  not  78 obviously  muscular a n t e r i o r p o r t i o n merging with the s l i g h t l y  broader, g r a n u l a r ,  posterior portion.  Oesophagus j o i n i n g  i n t e s t i n e without v a l v e ; j u n c t i o n u s u a l l y p o o r l y but  sometimes abrupt.  ilar,  the l e f t  Spicules  u s u a l l y pointed  subequal and s l i g h t l y and u s u a l l y longer  C u t i c l e moderately t h i c k , u s u a l l y with d e l i c a t e striations, irregular  demarcated,  than r i g h t . transverse  sometimes with bosses, sometimes with  annular r i d g e s .  dissim-  slightly  Vulva a n t e r i o r to end of oesophagus.  Caudal p a p i l l a e small and few i n number, i n v e n t r o l a t e r a l rows; preanal  p a p i l l a e o f t e n present.  digitiform. sent,  Amphids and two p a i r s * of c e p h a l i c p a p i l l a e pre-  o f t e n asymmetrically  developed.  T a i l of male s h o r t , not  arranged and sometimes  unequally  T a i l of one or both sexes o f t e n t e r m i n a t i n g  p a i r of f l e s h y protuberances. P a r a s i t e s of blood  vascular  M i c r o f i l a r i a e moderately  system ( u s u a l l y h e a r t ) ,  t i s s u e s , and serous c a v i t i e s of b i r d s .  B.  Comments on Recent Systems of C l a s s i f i c a t i o n of the Filarioidea filarial  short.  subcutane-  ous  The  ina  nematodes appear to be a h i g h l y  evolved  group of p a r a s i t e s which, i n the course of t h e i r s p e c i a l i z a t i o n as i n h a b i t a n t s aid  of t i s s u e s have l o s t many of the s t r u c t u r e s  i n the taxonomic s e p a r a t i o n  groups. strong  Structures buccal  of evident  capsules  of members of other  that  nematode  f u n c t i o n a l s i g n i f i c a n c e such as  ( i n forms which a t t a c h to the i n t e s t i n a l  *Dr. Anderson has informed the w r i t e r that S. alqonquinensis (Anderson, 1955) Anderson, 1961 possesses 2 p a i r s of asymmetric a l l y arranged p a p i l l a e , not 4 p a i r s as o r i g i n a l l y d e s c r i b e d ( 4 ) .  79 mucosa), or body spines  ( i n many p a r a s i t e s of the d i g e s t i v e ,  t r a c t ) , and such c h a r a c t e r s tion in  (cordons, c o l l a r s ,  the f i l a r i o i d s .  other  as e l a b o r a t e  c e p h a l i c ornamenta-  l i p s ) are r a t h e r c o n s i s t e n t l y l a c k i n g  Consequently, the f i l a r i a e resemble each  much more c l o s e l y than do many of the species  superfamilies,  and t h i s  t i v e characters  i n other  lack of r e a d i l y d i s c e r n i b l e separa-  has r e t a r d e d  the development of the s y s t e -  matics of the group. It lay  has long been known that some f i l a r i a l  embryonated eggs, whereas others  microfilariae  worms  produce embryos -  - which c i r c u l a t e i n the host's blood  or accu-  mulate i n the p e r i p h e r a l t i s s u e s .  Wehr (81), who appears to  have been the f i r s t  t h i s knowledge i n t o a  to incorporate  system of c l a s s i f i c a t i o n , r e c o g n i z e d filariidae, in  4 f a m i l i e s , Stephano-  Desmidocercidae, F i l a r i i d a e ,  1935, using  l a r v a l characters  and Dipetalonematidae,  t o d i s t i n g u i s h between the  l a s t two. In 1953 Chabaud and Choquet  (24) expanded and r e -  v i s e d Wehr's c l a s s i f i c a t i o n w i t h i n the same arrangement of families. 14)  Between 1955 and 1959, Anderson (4, 5, 7, 10, 13,  clarified  some of the r e l a t i o n s h i p s w i t h i n the F i l a r i i d a e ,  S t e p h a n o f i l a r i i d a e , Aproctinae, proposed and  and S p l e n d i d o f i l a r i i n a e , and  (7) a new f a m i l y D i p l o t r i a e n i d a e .  In 1959, Chabaud  Anderson (23.) r a t h e r e x t e n s i v e l y r e v i s e d the 1953 c l a s s i -  fication.  The l a t e s t  attempt  at a r e v i s i o n of the f i l a r i a l  nematodes i s that of Sonin i n 1963 (73). recent  Because these most  systems d i f f e r i n c e r t a i n b a s i c , and many minor p r e -  80 cepts, some d i s c u s s i o n of t h e i r warranted.  relative  merits seems  These systems can be summarized, t o subfamily  l e v e l , as f o l l o w s : Chabaud and Anderson 1959 Filarioidea  Sonin 1963 Filariata  Desmidocercidae  Aproctoidea Aproctidae  Diplotriaenidae Diplotriaeninae  Aproctinae  Dicheilonematinae  Tetracheilonematinae  Filariidae  Squamofilariinae  Filariinae  Splendidofilariidae  Tetracheilonematinae  Splendidofilariinae  Aproctinae  Eufilariinae Diplotriaenoidea  Setariidae  Diplotriaenidae  Stephanofilariinae  Diplotriaeninae  Setariinae  Dicheilonematinae  Onchocercid'ae  Lemdanidae  Oswaldofilariinae Icosiellinae  Lemdaninae  Dirofilariinae  Cardionematinae;  Onchocercinae  Icosiellinae  Splendidofilariinae  Oswa l d o f i l a r i i n a e  Ornithofilariinae*  Filarioidea Filariidae  Lemdaninae  Filariinae  Eufilariinae  Set a r i i d a e Setariinae Dirofilariinae Onchocercinae Stephanofilariinae * Suppressed  by Anderson 1961  (10).  81 The  authors of both of these systems of  c l a s s i f i c a t i o n s u b s c r i b e d to hypotheses worms are p o l y p h y l e t i c i n o r i g i n . gested that  "les f i l a i r e s  homogene, mais rassemblent detache's de d i f f e r e n t s  that the  filarial  Chabaud (22) had  sug-  ne r e p r e s e n t e n t pas un phylum un grand nombre de rameaux  S p i r u r i d e s a des periodes varie'es."  L a t e r , Anderson s p e c u l a t e d (6) that t h e l a z i i d - l i k e  ances-  t o r s gave r i s e t o the d i p e t a l o n e m a t i d s ( i . e . onchocercids) and "some of the f i l a r i i d s " ;  that  a i r sacs" ( i . e . d i p l o t r i a e n i d s )  " f i l a r i o i d s occurring i n  "may  r e p r e s e n t a separate  l i n e of e v o l u t i o n from that which gave r i s e to the dipetalonematids" (7) that  ( i . e . , o n c h o c e r c i d s ) , and,  "the S t e p h a n o f i l a r i i d a e "  to be the s t r o n g e s t evidence  ( i . e . S e t a r i i d a e ) "seem  f o r the b e l i e f that  f i l a r i o i d s have had a p o l y p h y l e t i c o r i g i n . of  subsequently  The  the absence  a i r sacs i n mammals seems to separate the S e t a r i i n a e  p h y l o g e n e t i c a l l y from the f i l a r i o i d s reptiles  of the a i r sacs of  and b i r d s with which they have a s u p e r f i c i a l r e -  semblance and with which they have h i t h e r t o classified."  (1958)  The major taxa i n the c l a s s i f i c a t i o n of  Chabaud and Anderson (23) are so arranged  as to i n d i c a t e  increasing biological  specialization,  (acquired a f t e r  they a s s e r t , confirms the  1953)  p o s i t i o n s of these taxa when based dence  been  (particularly  knowledge of whichrelative  on morphological  evi-  such f e a t u r e s as suggest i n c r e a s e d  r e d u c t i o n and s i m p l i f i c a t i o n  of p r i m i t i v e , i . e . , s p i r u r i d ,  82 structures).  Chabaud and Anderson i n c l u d e d the  Desmidocercidae,  which they regarded as intermediate  between the S p i r u r o i d e a and the F i l a r i o i d e a ,  i n the l a t t e r  s u p e r f a m i l y p a r t l y f o r the sake of convenience. in  a subsequent  paper  However,  (8) Anderson concluded that "'there  might be some j u s t i f i c a t i o n  f o r placing  .(the Desmido-  c e r c i d a e and the A p r o c t i n a e ) i n the same f a m i l y i n the f u t u r e e s p e c i a l l y as the Aproctinae are c l e a r l y u n r e l a t e d to  the F i l a r i i n a e with which they are p r e s e n t l y a s s o c i a t e d  in  the F i l a r i i d a e " .  The w r i t e r i s not convinced that  such  strong a f f i n i t y between the Aproctinae and the Desmidocercidae ter  e x i s t s , and would p r e f e r to leave the l a t -  f a m i l y r a t h e r l o o s e l y attached t o the F i l a r i o i d e a . Sonin  (73) expressed  f i l a r i a l worms can be arranged  the o p i n i o n that the i n three independent  groups  (Aproctoidea, D i p l o t r i a e n o i d e a , and F i l a r i o i d e a ) each of which has o r i g i n a t e d ancestors".  "at d i f f e r e n t times from  He c r i t i c i z e d  different  Chabaud and Anderson f o r p l a c i n g  undue emphasis on b i o l o g y and type of egg and l a r v a , and for  r e j e c t i n g the " c o r r e l a t i v e dependence" between s p i e - -  lar  s t r u c t u r e and c i r c u m o r a l e l e v a t i o n s proposed by  S k r j a b i n and considered by Sonin to be of primary tance f o r suprageneric s e p a r a t i o n . fallibility  of the system  c i t e d the genera  impor-  As an example of the  of Chabaud and Anderson, ,Sonin  Aprocta and S e r r a t o s p i c u l u m both of  which l a y t h i c k - s h e l l e d embryonated eggs, but which, as  83 a d u l t s , d i f f e r markedly and should not be considered c l o s e l y related.  Inasmuch as Chabaud and Anderson p l a c e d these  i n separate f a m i l i e s d i f f e r e n t i a t e d by c e p h a l i c and to a minor degree cluding spicules,  ornamentation,  by other m o r p h o l o g i c a l c h a r a c t e r s i n -  (but c e r t a i n l y not by t h e i r eggs and  which are very s i m i l a r ) , Sonin's c r i t i c i s m Sonin's  genera  seems to be  larvae, invalid.  c l a s s i f i c a t i o n embodies a number of e r r o r s  of omission and s e v e r a l n o m e n c l a t o r i a l d i s c r e p a n c i e s of which the f o l l o w i n g w i l l  serve as examples.  Monopetalonema D i e s i n g , 1861  l ) The  i s omitted e n t i r e l y .  e x p l a n a t i o n , Contortospiculum S k r j a b i n , 1915 f o r Dicheilonema 1915  genus  D i e s i n g , 1861,  2)  i s substituted  and S e r r a t o s p i c u l u m S k r j a b i n ,  i s named type genus of the D i c h e i l o n e m a t i n a e .  Cardionematinae  Yamaguti, 1941,  Without  3)  suppressed by Chabaud and  Anderson because i t s type and only s p e c i e s Cardionema ten Yamaguti, 1945  i s a metastrongyle,  Cardianema A l i c a t a , Onchocercidae  1933.  ( L e i p e r , 1911  i s resurrected for  4) For no apparent  reason  sub-fam.) i s r e p l a c e d by  S e t a r i i d a e S k r j a b i n and Schikhobalova,  1945  ( d e s p i t e the  a v a i l a b i l i t y of D i r o f i l a r i i d a e Sandground, 1921).  5) A d i a g -  n o s i s i s provided f o r the Lemdaninae but not f o r the Cardionematinae,  I c o s i e l l i n a e , or O s w a l d o f i l a r i i n a e  (? p o s s i b l y  because such diagnoses would c o n f l i c t with that of the f a m i l y , Lemdanidae). Filarioidea,  6) Sonin s t a t e s , i n h i s d i a g n o s i s of the " p a r a s i t e s of mammals e x c l u s i v e l y " thus  ignoring  the e x i s t e n c e of the s p e c i e s of S a u r o f i l a r i a C a b a l l e r o , 1945,  84 and  Macdonaldius Khanna, 1933  Oehoternella and  Caballero,  F o l e y e l l a Seurat,  1944  1917  (Onchocercinae) of r e p t i l e s , (Onchocercinae) of  (Dirofilariinae)  amphibians,  of r e p t i l e s  and  amphibians. Such e r r o r s and  misinterpretations  part r a t h e r p r e j u d i c e the reader  against  oniSonin's  accepting  he proposed.  Moreover, Sonin's hypothesis  o r i g i n of the  filarioids  the  of a t r i p h y l e t i c  i s not w e l l supported, p a r t i c u l a r l y  s i n c e i t i m p l i e s that m i c r o f i l a r i a e have a r i s e n i n 3 evolutionary  lines.  The  microfilarial  stage,  an  embryo, seems to have been i n t e r p o l a t e d i n t o the spiruroid  life  cycle.  i n p e r m i t t i n g the  I t s adaptive  separate  attenuated usual  s i g n i f i c a n c e i s apparent,  a d u l t s to occupy a great v a r i e t y of  deep w i t h i n the host's  t i s s u e s and,  Filarioidea  sites  at the same time, making  inoculum r e a d i l y a v a i l a b l e to haematophagous Throughout the  system  (Filariata  arthropods.  of Sonin), m i c r o f i l a r i a e  are e s s e n t i a l l y i d e n t i c a l i n morphology and  biology.  The  w r i t e r i s of the o p i n i o n that such a s t r i k i n g convergence is implicit  i n Sonin's hypothesis  lacks convincing  confirmation  i s u n l i k e l y , and  h i s hypothesis  as  since i t  should  be r e j e c -  ted. C e r t a i n l y , as Chabaud and the  1959  Anderson f r e e l y admit,-  system of c l a s s i f i c a t i o n embodied a number of uncer-  t a i n t i e s and  p r o v i s i o n a l conclusions.  very w e l l organized  Nevertheless,  i t is a  system, i n which the authors demonstrated  t h e i r f a m i l i a r i t y with most f i l a r i o i d  genera.  Until a  con-  85  siderable seem  volume  advisable  Anderson.  to  of  new  information  u t i l i z e  the  system  becomes  available,  proposed  by  i t  Chabaud  would and  86 PART TWO.  BIOLOGY  INTRODUCTION The f i r s t l i f e  c y c l e of a f i l a r i a l  worm, that of  Wuchereria b a n c r o f t i of man, was e l u c i d a t e d by Manson i n 1878. Since  then, the l i f e  viviparous  h i s t o r i e s of some 47 other  f i l a r i o i d s have been worked out; 35 of these belong  to the D i r o f i l a r i i n a e of  species of  and Onchocercinae, and most are p a r a s i t e s  mammals. Until recently, l i t t l e  the  transmission  of avian  a t t e n t i o n has been given to  f i l a r i o i d s , although as e a r l y as 1905  Dutton  (21) i n d i c a t e d that larvae he had found developing i n  biting  l i c e from A f r i c a n s w i f t s  those of F i l a r i a 1955,  (Cypselus  Gray) were  c v p s e l i Annett, Dutton and E l l i o t t , 1901. In  Raghavan and David  (44) noted that  l i s sum Chandler, 1929,* from an Indian Culex f a t i g a n s .  affinis  In 1956 Anderson  larvae of  Aproctoides  f r a n c o l i n , developed i n  (4) made a major c o n t r i b u t i o n  to the knowledge of s p l e n d i d o f i l a r i i n e b i o l o g y through h i s study of the development and t r a n s m i s s i o n fallisensis  (Anderson, 1954) of domestic ducks i n O n t a r i o .  Whereas Anderson found that black species  of S p i e n d i d o f i l a r i a  transmitted  flies  S. f a l l i s e n s i s .  sp., and E u f i l a r i a  of s e v e r a l  H i b l e r i n 1961, i n c r i m i n a t e d  b i t i n g midges ( C u l i c o i d e s ) as v e c t o r s "Chandlerella"  (Simuliidae)  f o r S p i e n d i d o f i l a r i a sp.,  sp. of Colorado magpies (32).  * This may be Pseudaproctoides p a p i l l a t u s ( A l i , 1957) Sonin, 1961.  87  In 1962, Nelson 1905  (39) r e p o r t e d  observations  biting  that he had v e r i f i e d  on the development of F i l a r i a  Dutton's  cypseli in a  louse, Dennvsus h i r u n d i s . A number of authors  have admitted  attempts to f i n d the v e c t o r s of v a r i o u s of b i r d s .  Gonnert  failure  i n their  splendidofilariines  (26), f o r i n s t a n c e , was unable t o d i s c o v e r  the v e c t o r s of S p l e n d i d o f i l a r i a mavis ( L e i p e r , 1909) by methods he d i d not d e s c r i b e .  In Georgia,  Robinson (47) allowed more  than 1000 mosquitoes of 13 species to feed on m i c r o f i l a r a e m i c crows, blue j a y s , white-throated  sparrows, and c a r d i n a l s i n a  s e r i e s of l a b o r a t o r y experiments, but found no s i g n i f i c a n t development i n any.  Odetoyinbo  (42), working with  which harbored C h a n d l e r e l l a g u i s c a l i to and  1904) was unable  o b t a i n development i n 4 species of l a b o r a t o r y - r e a r e d mosquitoes 2 species of b i r d mites.  B a s i c a l l y , these  because the experiments had ignored e p i z o o t i o l o g y of these attacked in  (Linstow,  grackles  the b i r d s i n t h e i r n a t u r a l h a b i t a t .  Anderson (3) u t i l i z e d  aspect  failed  of the  f i l a r i a s e s , v i z . , the arthropods  h i s i n v e s t i g a t i o n of the l i f e  Fallis  a vital  studies  which  By way of c o n t r a s t ,  c y c l e of S. f a l l i s e n s i s .  to advantage the knowledge acquired by  et .al. (25) that s i m u l i i d s , as v e c t o r s of the haemo-  s p o r i d i a n Leucocvtozoon simondi i n the same r e g i o n , f e d i n large numbers on domestic ducks. Bennett's study recovered  ( l l ) of the o r n i t h o p h i l i c  dipterans  from 20 species of b i r d s , exposed at v a r i o u s  heights  above ground i n d i f f e r e n t h a b i t a t s , has c o n t r i b u t e d much i n f o r m a t i o n on the h a b i t s of p o t e n t i a l f i l a r i a l  vectors.  Of  88 particular significance the b i t i n g  are h i s c o n c l u s i o n s that "....some of  f l i e s have marked host  These preferences mining the types  could c o n c e i v a b l y be as important of blood p a r a s i t e s found  of b i r d s as the s p e c i f i c i t y The was  and h a b i t a t p r e f e r e n c e s .  in certain  aim of the second p a r t of the present  filarial  worms are acquired by t h e i r t e t r a o n i d hosts, and then d i f f e r e n c e s i n prevalence  several f i l a r i a s e s  among s p e c i e s and  between geographic  r e g i o n s of the  With few filariine  exceptions,  s p e c i e s of host.  of avian f i l a r i o i d s  little  each species of s p l e n d i d o been r e p o r t e d  In most s t u d i e s on the b i o l o g y  or no attempt has been made to One  of the most  u b i q u i t o u s of North American s p l e n d i d o f i l a r i i n e s  found  (unpublished  Two  of the hosts f o r t h i s worm are  Bonasa umbellus  In view of t h i s ,  (5, 6) to Mexico (56), and  and  galli-  (5), and C o l i n u s v i r q i n i a n u s (18).  i n the l i g h t of Bennett's observations  haematophagous arthropods,  found  on  the w r i t e r deemed i t worthwhile to  e s t a b l i s h whether non-gallinaceous by the f i l a r i o i d s  the  i t i n a f u r t h e r 3 from B r i t i s h Columbia  data).  naceous b i r d s ,  is Aproctella  This p a r a s i t e has been r e p o r t e d i n 8  s p e c i e s of b i r d s from Ontario w r i t e r has  and  province.  d i s c o v e r i n d i c a t i o n s of host s p e c i f i c i t y .  s t o d d a r d i Cram, 1931.  to  of the  s t a t u s of host,  nematode d e s c r i b e d from b i r d s has  from only one  study  described  to d i s c o v e r the means by which these  account f o r the observed  species  of the p a r a s i t e s themselves."  to determine the i n c i d e n c e of the f i l a r i o i d s  i n Part One,  i n deter-  b i r d s as w e l l are  locally in tetraonids.  parasitized  89 In t h i s part of the t h e s i s new  data w i l l be  on the i n c i d e n c e of S k r i a b i n o c t a . f l e x i v a o i n a l i s . sp. B, and naceous and  galli-  b i r d s of s o u t h - c o a s t a l and  B r i t i s h Columbia, and M i c r o f i l a r i a and  Microfilaria  S p l e n d i d o f i l a r i a p e c t o r a l i s n. sp., i n the non-gallinaceous  presented  laoopodis  central  i n northern  B.  C.  Alaska. The  development of Sk.  sp. B i n t h e i r intermediate first  time,  with those  and  flexivaginalis  and  of  Mf.  hosts w i l l be d e s c r i b e d f o r the  s e v e r a l developmental f e a t u r e s w i l l be  compared  of other s p e c i e s . I t w i l l be shown that b i t i n g midges ( C u l i c o i d e s )  are the v e c t o r s of Sk.  f l e x i v a g i n a l i s . and black f l i e s  l i i d a e ) the v e c t o r s of Mf. of f l i e s  sp. B.  (Simu-  On the b a s i s of the numbers  c o l l e c t e d from e x p e r i m e n t a l l y exposed grouse the  activities  and  d e s c r i b e d , and  seasonal  abundance of these v e c t o r s w i l l  daily  be  the e f f e c t s of f a c t o r s of t h e i r environment  assessed. Finally,  the w r i t e r w i l l  knowledge a c q u i r e d i n t h i s i n f e c t i o n s , ecology  attempt to i n t e g r a t e the  study on i n c i d e n c e of  of avian hosts, and  ecology  filarial of v e c t o r s to  produce a comprehensive p i c t u r e of the e p i z o o t i o l o g y of flexivaginalis.  Mf.  sp. B,  i n the blue grouse and  and to a l e s s e r extent S.  other Tetraonidae  of B r i t i s h  Sk.  pectoralis.  Columbia.  90 MATERIALS 1. Study Areas Experimental  f i e l d r e s e a r c h was c a r r i e d out i n two  l o c a t i o n s , one near the Okanagan V a l l e y i n the i n t e r i o r of the p r o v i n c e , the other on Vancouver I s l a n d ( r e f e r r e d t o h e r e i n as "Trout Creek", and "Nanaimo Lakes", r e s p e c t i v e l y ) . of these  In both  areas c a p t i v e , i n f e c t e d blue grouse were exposed s y s t e -  m a t i c a l l y during the summer months i n order to d i s c o v e r the v e c t o r s of t h e i r f i l a r i o i d s , nent t o an understanding The  and t o acquire i n f o r m a t i o n p e r t i -  of the e p i z o o t i o l o g y of these  r e s e a r c h area u t i l i z e d  filariases.  i n the i n t e r i o r of the  p r o v i n c e i s i n the Trout Creek V a l l e y , some 20 m i l e s , by road, northwest of the south Okanagan town of West Summerland. r e s e a r c h cabin was on the v a l l e y f l o o r , beside  The  Trout Creek, at  an e l e v a t i o n of 3000 f e e t , about 1500 f e e t higher than Okanagan Lake.  In t h i s r e g i o n , Trout Creek i s a shallow  stream  between 10 and 30 f e e t wide, with a moderate to r a p i d  southeast  flow, and l i t t l e  aquatic v e g e t a t i o n .  of r i p a r i a n growth comprises  gravel-bottomed  A narrow zone  a l d e r , aspen, and r e d cedar.  For  the most p a r t , the f l o o d p l a i n i s very r e s t r i c t e d ,  nonexistent  i n some s t r e t c h e s , and i n others supporting stands  of aspen,  swampy a l d e r groves,  or wet or dry meadows.  The mountain  slopes r i s e with a moderate g r a d i e n t on both s i d e s of the narrow v a l l e y bottom, reaching an e l e v a t i o n 1200 f e e t above the r i v e r bed w i t h i n one-half to one m i l e .  Both slopes are i n -  c i s e d by numerous draws and g u l l e y s down which tumble small watercourses; most of these  are ephemeral and are dry by midsummer.  91 On the southeast  side of Trout Creek, the  Forest extends almost to streamside  and  comprises a  pine-Douglas f i r a s s o c i a t i o n with Engelmann spruce at  Sub-alpine lodgepole  appearing  higher e l e v a t i o n s . On  the northwest the lower l e v e l s of the  slopes  bear the t y p i c a l high Dry Forest B i o t i c Area a s s o c i a t i o n of yellow pine and Douglas f i r , with deciduous growth, aspen, i n the moist zone, and 500 lodgepole result  draws.  to 1000  This i s a r a t h e r narrow t r a n s i t i o n a l  f e e t above the r i v e r bed,  pine become i n c r e a s i n g l y e x t e n s i v e .  of past  stands  of  P a r t l y as a  l o g g i n g , grassy open benches and c l e a r i n g s  dotted with f i r s  are numerous.  P r e c i p i t a t i o n during the summer i s very temperatures r i s e 50-60°F at n i g h t .  to 80-95°F during the day, From l a t e May  swollen to many times snow m e l t i n g  mainly  and  slight;  drop to  to mid-June, Trout Creek i s  i t s subsequent volume by  at higher e l e v a t i o n s .  run-off.from  Throughout the summer,  water temperature remains below 60°F.  Many summer evenings  are accompanied by l i g h t to strong winds. Most exposures of grouse were c a r r i e d out on the northeast  slopes i n s e v e r a l mature Douglas f i r s ,  between 3800 and 4500 f e e t , i . e . , 800 stream  f e e t above the  bed. On Vancouver I s l a n d , the  of  to 1500  at e l e v a t i o n s  study  the North Nanaimo R i v e r , approximately  southwest of Nanaimo. gravel-bottomed southeast  and  The  stream with  area i s i n the 20 m i l e s , by  road,  North Nanaimo R i v e r i s a very little  aquatic v e g e t a t i o n .  valley  shallow, I t flows  enters the Nanaimo R i v e r j u s t east of the Nanaimo  92 Lakes, about 700  feet  above sea l e v e l .  Comparatively f l a t  t e r r a i n extends from one-half to t h r e e - q u a r t e r s of a mile on both s i d e s of the N. Nanaimo R i v e r to the foot of Mt. de Cosmos (4400 f e e t ) on the west, the east.  These  flats,  and B l a c k j a c k Ridge  once densely f o r e s t e d , have been  logged over and burned, and now  support r a t h e r  d i s t r i b u t e d r e g e n e r a t i n g Douglas which  (3000 feet) on  sparsely  f i r s up to 25 f e e t  tall  are most numerous along the banks of the former  grades.  Bordering some reaches of the r i v e r  p a r t s of the f l a t s ,  logging  and i n the moist  red a l d e r swales are dense  and  often  extensive. The s p r i n g f l o o d occurs i n A p r i l l e s s extreme  than t h a t of Trout Creek.  and May  and i s  Precipitation  during  the summer ranges from moderate to low; temperatures r i s e to 75-80°F  during the day and drop to 55-60°F  at n i g h t .  Strong  winds are r a r e . About  a mile north of F i r s t Lake  (the most e a s t e r l y  of the Nanaimo Lakes chain) a spur of i n t a c t c o n i f e r o u s  bush  extends n o r t h e a s t on a s l i g h t e l e v a t i o n to w i t h i n 200 yards of the North Nanaimo R i v e r . fir  At the edge of t h i s stand of Douglas  and western hemlock most of the exposures of grouse were  c a r r i e d out.  A dozen exposure  s i t e s were used at one time or  another, f a r t h e r i n from the stand, i n predominantly c o n i f e r o u s bush, near a small t r i b u t a r y of F i r s t  younger  Lake, on the  slopes of B l a c k j a c k Ridge, and i n deciduous areas, but it'was found that one p a r t i c u l a r Douglas  f i r at the edge of the spur  c o n s i s t e n t l y y i e l d e d l a r g e r numbers of o r n i t h o p h i l i c  dipterans.  9?.  This was  2.  exposure found  site  t o be  used  productive  f r e q u e n t l y i n 1961 again  i n 1964  and  1962,  and  (55).  D i s t r i b u t i o n , H a b i t s , and H a b i t a t P r e f e r e n c e of G a l l i n a c e o u s B i r d s i n B r i t i s h Columbia Ten  s p e c i e s of  America,  and  Alaska.  They are  of these,  2)  grouse,  Canachites  grouse;  ruffed  4)  ptarmigan, leucurus  Laqopus' l a q o p u s  (Montin);  (Bonaparte),  which  to-day  8)  was  chukar,  not  Franklin's (L.);  white-tailed  sage g r o u s e ,  subspecies  o f the  and  subspecific  and  habitat preference  37,  53)  5)  rock  ptarmigan, Pedioecetes  Centrocercus  urophasianus  infrequent occurrence  considered  British  i n the  Columbia  p a r t r i d g e , and  present  but  California  in study.  phasianids, only  four,  quail  remain  numbers.  Appendix VII the  and  spruce  s h a r p - t a i l e d grouse,  of v e r y  into  gray  i n appreciable  and  3)  s p e c i e s of G a l l i f o r m e s , m o s t l y  have been i n t r o d u c e d pheasant,  7)  i s now  and  6)  North  obscurus  !  including  in  Columbia  Dendraqapus  ptarmigan,  ( L . ) ; and  present  Bonasa u m b e l l u s ( L . ) ;  (Richardson);  Eight  grouse,  (L.),  mutus  Columbia  in British  canadensis  phasianellus  British  eight occur  grouse,  willow L.  t e t r a o n i d b i r d s are  l ) blue  (Say);  L.  was  lists  the  names and  distribution  f o u r common t e t r a o n i d s of  names u s e d h e r e are  from p e r s o n a l  and  B.  i n f o r m a t i o n on  from v a r i o u s  sources  C.  for  Specific  distribution  ( l , 2, 17,  27,  observation.  * A c c o r d i n g t o G u i g e t ( 2 7 ) , c a p e r c a i l l i e and w i l d t u r k e y , i n t r o d u c e d i n t h e p a s t , have d i e d o u t , and m o u n t a i n q u a i l are p r e s e n t i n low numbers on s o u t h e r n V a n c o u v e r I s l a n d . I n 1947 (37) a few b o b - w h i t e q u a i l were t h o u g h t to have s u r v i v e d n e a r H u n t i n g d o n .  Although  s e v e r a l s p e c i e s of t e t r a o n i d s c a r r y out  v e r t i c a l m i g r a t i o n s , u s u a l l y i n the s p r i n g and f a l l , year-round areas.  r e s i d e n t s i n the general v i c i n i t y  breeding  This c h a r a c t e r i s t i c has c o n s i d e r a b l e bearing on the  e p i z o o t i o l o g y of the f i l a r i a l i m p l i e s that these locality Blue  of t h e i r  a l l are  i n f e c t i o n s encountered,  for i t  i n f e c t i o n s were acquired very c l o s e to the  i n which the b i r d s were c o l l e c t e d .  grouse Throughout much of B r i t i s h Columbia blue grouse  are the most abundant g a l l i f o r m . (37)  "at one season or another  According  to Munro and Cowan  (they frequent)  from sea l e v e l to the alplands where these  a l l biotic  adjoin."  however, some regions not occupied by blue grouse,  areas  There are,  the most  notable of these being the r o l l i n g • p a r k l a n d s known as the "Cariboo" and the " C h i l c o t i n " .  In g e n e r a l , blue grouse  c o n i f e r o u s f o r e s t s , p a r t i c u l a r l y those  inhabit  i n which Douglas f i r  predominates, at moderate e l e v a t i o n s . Large  areas of Vancouver I s l a n d , d e f o r e s t e d by  logging and widespread f i r e s , able f o r dramatic These burnt-over  have proved  e x c e p t i o n a l l y favour-  i n c r e a s e s i n the numbers of D. o. f u l i g i n o s u s r e g i o n s c o n s t i t u t e d the breeding grounds f o r  the bulk of the Vancouver I s l a n d blue grouse i n r e c e n t years. Subsequent s u c c e s s i o n a l changes i n these r e g e n e r a t i o n to deciduous able f o r continued these  support  areas,  cover, have rendered  involving them l e s s  favour  of high p o p u l a t i o n s and i n most of  areas blue grouse are d e c l i n i n g i n abundance.  In t h i s  h a b i t a t the a c t i v i t i e s of the males as w e l l as the hens and  95 chicks  are mainly t e r r e s t r i a l .  interior,  and  populations  In the  seem to be  low  and  s t a b l e , and  blue grouse, D.  o.  follow a pattern In both regions  well  9  there,  more of  the  arboreal.  Both on Vancouver I s l a n d  but  the  i n v i r g i n timber on Vancouver I s l a n d , blue grouse  a c t i v i t i e s of these b i r d s are  vations,  open f o r e s t s of  f u l i q i n o s u s and  and  i n the Okanagan r e g i o n ,  o,  D.  pallidus. respectively,  of seasonal movement to d i f f e r e n t a l t i t u d e s . most blue grouse winter at f a i r l y high  some can  be  found there  throughout the  ele-  summer as  perhaps comprising b i r d s that breed near a l p i n e meadows  and  at t i m b e r l i n e ,  the  y e a r l i n g grouse as w e l l  personal  and  many y e a r l i n g males. as the  communication, 1965)  adults  descend the  In March most of  (Dr.  J. F.  slopes  benchlands, n a t u r a l or man-made c l e a r i n g s and, I s l a n d , to the cover by  p l a i n s and  logging  and  Bendell,  to more open  on Vancouver  v a l l e y bottoms denuded of f o r e s t  fire.  In f o r e s t e d c o a s t a l areas male blue grouse, D. fuliqinosus, emitting  appear to e s t a b l i s h p r i m a r i l y a r b o r e a l  t h e i r v e n t r i l o q u i a l "hooting"  from the  (20  to 50  the  burnt-over areas of Vancouver I s l a n d  perforce,  o.  territories,  lower branches  f e e t above the ground) of s e l e c t e d Douglas f i r s .  terrestrial,  u s u a l l y i n the  "hooting"  sites  s h e l t e r of f a l l e n  In  are, trees  or beneath a Douglas f i r . P a r t i c u l a r l y favoured are-clumps of small  firs  on the  elevated  In the w r i t e r ' s the  borders of logging  and  experience, cocks of D.  fire  roads.  o.  pallidus.  i n t e r i o r blue grouse, i n v a r i a b l y "hoot" from the  When d i s t u r b e d ,  ground.  grouse of t h i s race tend to take s h e l t e r  initially  96  among the branches of a v a i l a b l e c o n i f e r s , u s u a l l y Douglas from which they are o f t e n d i f f i c u l t Nesting s i t e s courses or moist  to f l u s h .  are u s u a l l y not f a r from small water-  d e p r e s s i o n s , o f t e n i n exposed l o c a t i o n s - w i t h i n  c l e a r i n g s , on the open "burns", of  firs,  or near the s p a r s e l y t r e e d edges  stands of Douglas f i r , yellow pine or lodgepole pine, but  are not i n f r e q u e n t l y i n moderately  dense bush.  Peak hatching  times range from e a r l y to mid-June on Vancouver I s l a n d ; few broods are brought o f f before the end of May, and the w r i t e r has  encountered  c h i c k s at Cowichan, Nanaimo Lakes, and Quinsam  that must have hatched of  near the end of J u l y .  the province hatching occurs  In the i n t e r i o r  slightly earlier.  On the study  area i n the Trout Creek V a l l e y , 20 miles west of West Summerland, the time  of peak hatching was about June 1; however, on the  hot, dry benchlands of the northern Okanagan, near Oyama and Kelowna, most c h i c k s leave the egg a week to ten days e a r l i e r , and  some by the middle During  of May.  their first  2 weeks of l i f e  the c h i c k s are  virtually  incapable of f l i g h t ,  and r e q u i r e frequent brooding by  the hen.  They appear t o begin f o r a g i n g seldom before  morning s u n l i g h t has e l e v a t e d a i r temperatures the dew.  P r e f e r r e d feeding areas  direct  and evaporated  seem to be open grassy meadows  supporting low v e g e t a t i o n . By mid-July  about h a l f the adult males of D. o.  f u l i q i n o s u s have departed  * The w r i t e r c o l l e c t e d Kelowna.  from the breeding grounds f o r the  a c l u t c h , pipped on May 14, 1960, from  97  h i g h e r e l e v a t i o n of nearby r i d g e s and mountainsides, and, u s u a l l y , by the end of August only the hens and t h e i r almost f u l l y grown broods, o f t e n c o n s i d e r a b l y aggregated, remain on the flats  and lower slopes of Vancouver  Island.  By mid-October  the  summer range has been completely d e s e r t e d . In c o n t r a s t , the i n t e r i o r blue grouse, D. o.  pallidus.  which breeds w e l l above the v a l l e y f l o o r , tends to descend i n l a t e J u l yi. and e a r l y A u g u s t , . p o s s i b l y i n search of water more verdant v e g e t a t i o n .  and  At t h i s time,, much of the v e g e t a t i o n  has been d e s s i c a t e d by s e v e r a l months of high temperatures  and  almost continuous sunshine, and most of the small watercourses have d r i e d up.  Many cocks are present with the hens and young  but i t appears that some adult males migrate upwards during August.  Many b i r d s are s t i l l  to be found at low e l e v a t i o n s i n  September, but probably by the f i r s t  s n o w f a l l most have reached  t h e i r w i n t e r i n g ranges at h i g h e r a l t i t u d e s . Ruffed  grouse U n l i k e blue grouse, r u f f e d grouse e x h i b i t no ten-  dency  to v e r t i c a l m i g r a t i o n .  Throughout  to be found i n deciduous and mixed r i p a r i a n growth,  the year they are  bush, p a r t i c u l a r l y i n  i n the v a l l e y bottoms.  With the  development  of e x t e n s i v e a l d e r swales on the burnt-over areas of Vancouver I s l a n d , r u f f e d grouse have become i n c r e a s i n g l y abundant blue grouse p o p u l a t i o n s have"waned. the slopes or r i d g e s of Vancouver  While seldom found on  I s l a n d , they are not uncommonly  encountered at moderate e l e v a t i o n s i n the i n t e r i o r , where aspen abound,  as  especially  98 On Vancouver I s l a n d , r u f f e d grouse c h i c k s hatch slightly May.  earlier  than blue grouse,  In the i n t e r i o r ,  u s u a l l y about the end of  at l e a s t i n Trout Creek' V a l l e y , most  broods appear i n the second week of June. Ruffed grouse occur throughout the northern coast) deciduous  the province  (except  and as f a r north as c e n t r a l Alaska, i n  and mixed f o r e s t a s s o c i a t i o n s .  Spruce grouse This species occurs north as f a r as northern and  i s g e n e r a l l y r e s t r i c t e d to sub-alpine and b o r e a l  forests,  although  i t s southern  grouse i s o f t e n found  Alaska,  spruce  counterpart, the F r a n k l i n ' s  at lower e l e v a t i o n s , u s u a l l y i n lodgepole  p i n e , and not i n f r e q u e n t l y i n areas occupied  by blue and r u f f e d  grouse. Ptarmigan Willow  ptarmigan,  Laqopus lagopus.  Laqopus mutus. are circumboreal  ptarmigan,  s p e c i e s o c c u r r i n g from e a s t e r n  Europe across northern A s i a and northern North t a i l e d ptarmigan,  and rock  America.  White-  Laqopus l e u c u r u s . on the other hand,:is  con-  f i n e d to western North  America, from c e n t r a l Alaska to northern  New Mexico.  species breeds from the tundra and c o a s t a l  The-' f i r s t  f l a t s i n the extreme north t o moist  plateaus at t i m b e r l i n e and  high a l p i n e meadows f a r t h e r south, whereas rock ptarmigan s p a r s e l y vegetated line.  rocky a l p i n e barrens, o f t e n f a r above  selects timber-  W h i t e - t a i l e d ptarmigan are r e s t r i c t e d to the a l p i n e  meadows and p l a t e a u s near or above t i m b e r l i n e where, according to Weeden (53) and Choate  (17), they p r e f e r moist  outcrops or  99 ledges which provide rocky cover and i n the  form of low  plants..  ptarmigan o v e r l a p s , occupy, i n that Sharp-tailed  order,  3  elevations.  grouse  are  but  i t s centre  of abundance i n the  C h i l c o t i n regions  interrupted- by mixed and  southern part  proximity,  d i s t r i b u t i o n of the  summer range at i n c r e a s i n g  lands of the Cariboo and  In the  Where the  vegetation  willow, rock, and w h i t e - t a i l e d ptarmigan  This grouse has  grasslands  which bear ample  where the  coniferous  park-  rolling  forests  of i t s range blue grouse occur i n close  f a r t h e r north,  where t h i s species  s h a r p - t a i l e d grouse come i n t o contact  i s absent,  with F r a n k l i n ' s  and  ruffed  grouse. Phasianids C a l i f o r n i a q u a i l , Lophortvx c a l i f o r n i c u s . are locally  abundant on southern Vancouver I s l a n d  southern Okanagan V a l l e y . and  established  overlap  become success-  on southern Vancouver I s l a n d , the  mainland, along the Valley.  to  galliform birds.  Pheasant, Phasianus c o l c h i c u s . has fully  i n the  Since they occur close to s e t t l e d  farming areas t h e i r l o c a l d i s t r i b u t i o n f a i l s  that of n a t i v e  it  and  Thompson R i v e r ,  and  lower  throughout the Okanagan  It occupies c u l t i v a t e d land f o r the most part which  shares, o c c a s i o n a l l y , with s h a r p - t a i l e d grouse and  gray  partridge» Gray p a r t r i d g e , Perdix Thompson River V a l l e y and  the  perdix,  i s present i n the  Fras.er Canyon, l e s s abundantly  100 elsewhere, and occurs on a g r i c u l t u r a l land and on some of the lower benches  of the v a l l e y s .  Chukar,  A l e c t o r i s graeca. has proved a s u c c e s s f u l  i n t r o d u c t i o n i n many of the a r i d areas of the southern Cariboo where other g a l l i n a c e o u s b i r d s are seldom  found.  3. I n f e c t i o n s i n Grouse Used f o r Exposure Blue grouse captured on Vancouver  I s l a n d or i n  s o u t h - c e n t r a l B r i t i s h Columbia were exposed r o u t i n e l y i n both study areas to a t t r a c t b i t i n g  insects.  Pertinent  biographical  i n f o r m a t i o n f o r these grouse i s as f o l l o w s : #9 - adult female D,.o. p a l l i d u s . captured near P e n t i c t o n 15/5/59, exposed of a s p e r g i l l o s i s .  1959 at Trout Creek, died  25/4/61  Harbored Mf. f l e x i v a q i n a l i s . and Mf. sp. B  or Mf, p e c t o r a l i s throughout c a p t i v i t y .  No adults found at  necropsy. #49 - adult female D. o. p a l l i d u s . captured near Kelowna 14/5/60, exposed  1960 at Trout Creek and 1961 at  Nanaimo Lakes, died 30/7/61 of (?) e r y t h r o c y t o p e n i a . Mf. sp. B or Mf. p e c t o r a l i s throughout c a p t i v i t y . of p e c t o r a l y e l l o w i n g when captured.  Harbored  No evidence  No a d u l t s found at  necropsy. #15 - adult male D. o. p a l l i d u s . captured near Oyama 4/8/60, exposed  1960 Trout Creek and 1962 Nanaimo Lakes,  d i e d 13/9/62 of a s p e r g i l l o s i s .  Harbored Mf. f l e x i v a q i n a l i s .  and Mf. sp. B. or Mf. p e c t o r a l i s throughout c a p t i v i t y . evidence of p e c t o r a l y e l l o w i n g .  No adults found.  No  101 #24 - adult female D. o. f u l i o i n o s u s .  captured at  I-  Quinsam 7/8/59, exposed aspergillosis.  1961 Nanaimo Lakes, died 27/10/61 of  Harbored Mf. sp. B throughout c a p t i v i t y .  No  adult's found. #25 - adult female D. o. fulioinosus« Quinsam 7/8/59, exposed  captured at  1960 Trout Creek and Nanaimo Lakes,  died 6/6/61 of a s p e r g i l l o s i s .  Harbored Mf.  and Mf. sp. B throughout c a p t i v i t y .  flexivaoinalis  No adults  found.  #3 - adult female D. o. f u l i o i n o s u s . Nanaimo Lakes 23/6/61, exposed of  (?) m a l n u t r i t i o n .  flexivaginalis.  1961 Nanaimo Lakes, died 3/7/61  Harbored very heavy / i n f e c t i o n  Y i e l d e d 7 male and 10 female S.  #4 - adult female D. o. f u l i o i n o s u s . Quinsam 27/7/61, exposed f i c e d 25/1/63.  captured at  flexivaginalis. captured at  1961 and 1962 Nanaimo Lakes, s a c r i -  Harbored Mf. sp. B.  No adults  found.  #33 - y e a r l i n g female D. o. f u l i o i n o s u s . at Quinsam 11/6/62, exposed 27/1/63.  1962 Nanaimo Lakes,  Harbored Mf. sp. B.  No adults  Nanaimo Lakes 19/6/62, exposed Harbored very heavy  8/3/64.  sacrificed  captured at  1962 Nanaimo Lakes, escaped infection  #8 - adult female D. o. Nanaimo Lakes 21/6/62, exposed  captured  found.  #7 - adult female D. o. f u l i o i n o s u s .  21/6/62.  of Mf.  of Mf.• f l e x i v a g i n a l i s .  fulioinosus.  captured at  1962 Nanaimo Lakes,  Harbored heavy i n f e c t i o n  sacrificed  of Mf. f l e x i v a g i n a l i s .  Y i e l d e d 7 male and 6 female S k . f l e x i v a g i n a l i s .  102 METHODS 1. C o l l e c t i o n of Data on N a t u r a l I n f e c t i o n s With a view to a s s e s s i n g d i s t r i b u t i o n , of i n f e c t i o n ,  and apparent  of the Tetraonidae  host  specificity  intensity  of the f i l a r i o i d s  of B r i t i s h Columbia, the w r i t e r examined  as r e p r e s e n t a t i v e a sample of these b i r d s as p o s s i b l e . Information  on n a t u r a l i n f e c t i o n s was obtained  by the w r i t e r and by others who c o l l e c t e d g a l l i n a c e o u s b i r d s , blood smears, or adult f i l a r i a e  from v a r i o u s r e g i o n s of  B r i t i s h Columbia and Alaska. E a r l y i n the study i t was d i s c o v e r e d that when p e r i p h e r a l blood seemed negative, m i c r o f i l a r a e m i a s could o f t e n be detected i n a drop of blood from the lung. quently, blood smears were made from t h i s organ  Conse-  throughout  most of the study. In order to determine whether the f i l a r i o i d s  found  i n t e t r a o n i d s p a r a s i t i z e d other b i r d s as w e l l , a t o t a l of 225  adult non-gallinaceous  was  examined on or near grouse range, mainly  and Nanaimo Lakes.  b i r d s r e p r e s e n t i n g 27 f a m i l i e s at Trout Creek  Despite p a i n s t a k i n g search adult  were very seldom recovered  filarioids  from m i c r o f i l a r a e m i c non-gallinaceous  birds. 2. Determination  of M i c r o f i l a r i a l  Periodicity  To d i s c o v e r whether Mf. sp. B, and Mf, f l e x i v a q i n a l i s e x h i b i t rhythmic  f l u c t u a t i o n s i n numbers i n p e r i p h e r a l blood,  a c t u a l counts were made at 2- or 3-hour i n t e r v a l s , f o r 24 or  103 36 hours,  as f o l l o w s :  b r a c h i a l v e i n was capillary pipette  Blood w e l l i n g from acupuncture  taken i n t o a h e p a r i n - r i n s e d , ' s t a n d a r d i z e d 9  d e l i v e r e d onto a 50 by 75 mm  and q u i c k l y covered with a 22 by- 50 mm lightly  glass slide  v a s e l i n e - r i n g e d cover-  s.lip which was  then tapped  This mount was  then secured on another 50 by 75 mm  to spread  v i o u s l y r u l e d i n a g r i d , each square s m a l l e r than the diameter a dissecting  the blood  of which was  of the high-power f i e l d  evenly.  slide  pre-  slightly (x30)  of  microscope.  One  of the .grouse t e s t e d (#24)  harbored  only  sp. B, the other was  i n f e c t e d with both Mf;  flexivaqinalis.  of the f o r e g o i n g method d i d not  Use  accurate d i f f e r e n t i a t i o n of the two to  of a  species.  sp. B and  In an  e s t a b l i s h the numbers of each s p e c i e s present  Mf. Mf.  permit attempt  at the  times  t e s t e d , the contents of the c a p i l l a r y p i p e t t e were d e l i v e r e d onto s l i d e s stained  and made i n t o smears'.  These were l a t e r  fixed,  and examined.  3. Maintenance of Captive Grouse Adult blue grouse were captured by a pole-and-noose technique developed  by Dr. J . F. B e n d e l l ; by c l a p p i n g a long-  handled, wide-mouthed landing net over them as they  incubated  c l u t c h e s ; or by l u r i n g them within, n e t t i n g range with simulated or  actual  " d i s t r e s s - c a l l s " \ o f grouse c h i c k s . Grouse c h i c k s were captured by hand, net, or noose.  As w e l l , three c l u t c h e s were incubated s u c c e s s f u l l y a f t e r t r a n s port, from the f i e l d warmed g r a i n .  to the l a b o r a t o r y i n i n s u l a t e d jugs of  104 In the l a b o r a t o r y , c h i c k s were reared i n an e l e c t r i c brooder; i n the f i e l d was used.  a converted, kerosene-heated i n c u b a t o r  As soon as p o s s i b l e a f t e r hatching or capture  chicks were induced to eat moistened, commercial mash by combining i t with chopped  fresh f r u i t .  were f e d turkey grower or turkey poult with No. 2 g r a n i t e g r i t  added.  turkey-starter Older grouse  "krumbles" ad, l i b i t u m  This d i e t was  supplemented  f r e q u e n t l y with leaves of hare's ear (Hypochaeris s p . ) , c l o v e r , and d a n d e l i o n . When about one month of age the c h i c k s were t r a n s f e r r e d to cages of g a l v a n i z e d , welded-mesh wire. (17-24 by 15 by 16 inches) were suspended  These  cages  from "Dexion" frames  indoors. Despite a few deaths due t o a c c i d e n t s and f a i l u r e to eat, m o r t a l i t y of chicks was c o n s i s t e n t l y low.  That of y e a r l i n g s  and a d u l t s was somewhat higher, the p r i n c i p a l causes of death being a s p e r g i l l o s i s a more d e t a i l e d  and an undetermined g a s t r o - e n t e r i t i s .  account of the techniques employed  For  i n incubating,  r e a r i n g , and m a i n t a i n i n g these c a p t i v e blue grouse the reader i s r e f e r r e d to Jensen 1962 (35). 4. Exposure of I n f e c t e d Grouse Biting Flies  and C o l l e c t i o n .of  Captive blue grouse harboring m i c r o f i l a r i a e were exposed to n a t u r a l p o p u l a t i o n s of b i t i n g f l i e s separated l o c a l i t i e s abundant  at two widely  i n B. C. where grouse were known to be  and known to have f i l a r i a l  infections.  Throughout  the summer of 1961, and during June 1962 exposures were made about 700 feet  above sea l e v e l at Nanaimo Lakes.  105 D a i l y exposures i n the i n t e r i o r of the province during the summer of 1959 and on Vancouver I s l a n d i n 1961 provided data on the seasonal v a r i a t i o n i n p o p u l a t i o n s of grouse-biting f l i e s .  Records of temperature  c o n d i t i o n s were made at exposure times. at  and other weather  Grouse were exposed  i n t e r v a l s throughout the day to determine peak b i t i n g  times, and at v a r i o u s s i t e s , both oh-..the grcound and i n t r e e s to  o b t a i n some knowledge of the d i s t r i b u t i o n of p o t e n t i a l  vectors. In  1958 the w r i t e r ' s observations suggested that  blue grouse c h i c k s i n the Okanagan r e g i o n became i n f e c t e d before they were o l d enough to perch i n t r e e s .  In  1959,  because of the i n d i c a t i o n s of t r a n s m i s s i o n near ground an i n f e c t e d grouse was l a t e May  exposed mainly on the ground from  to e a r l y J u l y .  c o n s i s t e n t l y negative.  Results of these exposures were I t was  then found that  exposures were much more p r o d u c t i v e of f l i e s . for  level  arboreal Consequently,  the remainder of the study most of the exposures were  c a r r i e d out i n t r e e s branches 35-50 f e e t  (Douglas f i r s ) , u s u a l l y beside l i v i n g above ground.  P r i o r to exposure, a hood of black c l o t h with a dome-fastener at the t h r o a t and an opening f o r the beak was placed over the head of each grouse to be used, because unhooded, they f l u t t e r e d  and paced almost c o n s t a n t l y .  cages were 15-inch cubes of 1.5 l i f t - r o p e was  Exposure  i n c h wire-mesh to which the  attached by a swivel-mounted  s n a p - c l i p which  e f f e c t i v e l y prevented the cage from s p i n n i n g and ensured  106 r a p i d disengagement  from the rope when r e q u i r e d .  more e l a b o r a t e methods f o r r a i s i n g u n s a t i s f a c t o r y , i t was found that  After  several  and lowering the cage proved a length of No. 7 hard-  surface sash cord simply passed over the branch was i d e a l , f o r the weight of the caged grouse was j u s t keep the rope running smoothly.  s u f f i c i e n t to  The caged grouse was h o i s t e d  to b r a n c h - l e v e l i n 10-15 seconds, the rope was anchored, and the  bird  left  suspended  f o r 10-15 minutes.  The remainder  of the exposure method, and techniques f o r c o l l e c t i n g flies  have been d e s c r i b e d elsewhere  engorged ;  (11, 55).  On s e v e r a l occasions large numbers of mosquitoes hovering about man were captured and placed under the c o l l e c t i n g cage with a grouse. On the evening of J u l y 29, 196L, many louse  flies  (Hippoboscidae) were noted i n the a i r . S e v e r a l dozen of thesewere captured with a net and l e f t overnight with a hooded grouse.  under the c o l l e c t i n g  Attempts t o o b t a i n hippo-  boscids with grouse exposed during d a y l i g h t  hours,were  u n s u c c e s s f u l because the high temperatures r e s u l t i n g repeated or prolonged r e t e n t i o n w i t h i n the c o l l e c t i n g were very d e b i l i t a t i n g 5. Maintenance  from cage  to the grouse.  of Engorged  Engorged  cage  flies  Flies  that had been captured from exposed  grouse were maintained i n cages l i k e those developed by Anderson the  ( 4 ) . S e v e r a l l a y e r s of paper t o w e l l i n g placed on  bottom of the cage and secured to the s i d e s with masking  tape were moistened thoroughly with b o i l e d water each day to  107 provide high'humidity.  To reduce' evaporation and exclude  light  a cardboard  d i s c was placed over the armhole and a cardboard  cover taped  over the top and f r o n t of the cage.  was  The cover  removed f o r s e v e r a l hours d a i l y to allow the f l i e s  t c feed.  Anderson's method (4) of feeding by p r o v i d i n g bags of dry sucrose  and cheesecloth-covered  fication.  j a r s of water r e q u i r e d modi-  The j a r s of water made i t d i f f i c u l t  to move the  cages,  the sucrose  a t t r a c t e d carpenter ants which invaded the  cages,  and there was no i n d i c a t i o n that the f l i e s  i n g e s t e d the food or water. nique  proved  covered, was  successful.  actually  E v e n t u a l l y the f o l l o w i n g tech-  Each day, when the cages were un-  a viscous, s o l u t i o n of l i q u i d honey i n b o i l e d water  painted i n a narrow s t r i p  Culicoides  across the nylon screen.  and s i m u l i i d s consumed t h i s  hesitation.  s o l u t i o n with  Both  little  Before c o v e r i n g the cages the screen was g e n t l y  washed with a stream  of b o i l e d water to remove the syrup.  S u r v i v a l was g r e a t l y improved a f t e r t h i s method of feeding was  introduced..  When s a c r i f i c e d ,  some s i m u l i i d s had s u r v i v e d  40 days a f t e r capture, C u l i c o i d e s up to 43 days and Mansonia up t o 18 days.  S u r v i v a l might be f u r t h e r enhanced i n smaller,  cages with p a r t i a l l y  solid  f r o n t s , and by the use of a mould-  inhibitor. 6. Developmental  Stages  In.sects which had fed. on blue grouse i n f e c t e d w i t h Sk.  f l e x i v a q i n a l i s and/or Mf. sp. B were d i s s e c t e d at i n t e r v a l s  in physiological saline  and examined f o r developing l a r v a e .  A n a e s t h e t i z a t i o n of f l i e s with e t h y l acetate  facilitated  108 d i s s e c t i o n and had no n o t i c e a b l e e f f e c t on the a c t i v i t y of larvae.  Whenever p o s s i b l e , l a r v a e were s t u d i e d a l i v e , i n  s a l i n e , under a v a s e l i n e - r i n g e d c o v e r s l i p .  Others,  preserved  i n 70% ethanol or removed from preserved f l i e s , were t r a n s f e r r e d to  t% g l y c e r i n e - a l c o h o l , and examined i n g l y c e r i n e mounts a f t e r  the a l c o h o l had been allowed to evaporate  slowly.  Some l a r v a e  were s t a i n e d with c o t t o n blue i n l a c t o p h e n o l , but unstained l a r v a e , p a r t i c u l a r l y when a l i v e , were the most for  satisfactory  study. Whenever f l i e s were being maintained  a constantly  r e c o r d i n g thermograph was i n o p e r a t i o n beside the f l y cages. Readings at 3-hour i n t e r v a l s were used temperatures  to c a l c u l a t e the mean  f o r the p e r i o d s of l a r v a l development.  7. Experimental In  Transmission  the s p r i n g of 1959 i t was p o s s i b l e to capture  a l i v e only one m i c r o f i l a r a e m i c grouse p a l l i d u s #9); i t harbored and what were thought  (Dendraqapus  the m i c r o f i l a r i a e  obscurus  of Sk. f l e x i v a g i n a l i s  to be those of S. p e c t o r a l i s .  was exposed at Trout Creek, and by the time  suitable  This b i r d inter-  mediate hosts had been c o l l e c t e d i t had been found that capture of  other i n f e c t e d grouse  difficult.  Consequently,  i n t h i s r e g i o n would be p r o h i b i t i v e l y i t was decided to use the larvae  developing i n these f l i e s t o i n f e c t young grouse for  ( r a t h e r than  a study of l a r v a l development) with the i n t e n t i o n of producing  heavy i n f e c t i o n s i n as many semi-domesticated possible.  I t was envisaged  n a t u r a l hosts as  that besides p r o v i d i n g i n f o r m a t i o n  on a e t i o l o g y these b i r d s would be exposed the f o l l o w i n g summer to  o b t a i n developing larvae of S. p e c t o r a l i s .  109 Larvae of what was removed that  into physiological  thought to be S. p e c t o r a l i s were  saline  had fed on grouse #9 between  larvae  from the heads of s i m u l i i d s 9 and 17 days e a r l i e r .  were drawn i n t o a hypodermic syringe  equipped with a  No.  19 needle and i n j e c t e d under the p e c t o r a l  the  brachial vein  s k i n or i n t o  of 9 young grouse which had been reared  i n c a p t i v i t y from eggs or very young c h i c k s .  The  blood of a l l r e c i p i e n t grouse had been examined  peripheral  and found  negative f o r m i c r o f i l a r i a e and other blood p a r a s i t e s the  following  'These  inoculations  before-  were c a r r i e d out:  1) J u l y 30 - 40 l a r v a e , J u l y 31 - 110 l a r v a e , subcutaneously i n t o 70-day o l d female Dendragapus obscurus pallidus. 2) J u l y 31 - 97 l a r v a e , August 2 - 104 l a r v a e , subcutaneously i n t o 60-day o l d male Bonasa u. a f f i n i s . 70-day  3) August 3 - 9 4 larvae, o l d female D. o.. p a l l i d u s .  subcutaneously i n t o  4) August 12 - 81 l a r v a e , August 15 - 176 subcutaneously i n t o 70-day o l d male D. o, p a l l i d u s .  larvae,  5) August 16 - 97 l a r v a e , August 17 - 77 l a r v a e , subcutaneously i n t o 70-day o l d female D.. o. p a l l i d u s . • 77-day  6) August 18 - 83 l a r v a e , o l d female D. o. p a l l i d u s .  subcutaneously i n t o  80-day  l) August 23 - 94 l a r v a e , o l d male D. o. p a l l i d u s .  subcutaneously i n t o  8) August 24 - 129 l a r v a e , o l d female D. p_. p a l l i d u s .  intravenously  into  9) August 24 - 119 l a r v a e , y e a r l i n g male D. o. f u l i g i n o s u s .  intravenously  into  100-day  In a d d i t i o n , from a f r e s h l y k i l l e d the  pectoral  about 25 subadult S. p e c t o r a l i s  removed  blue grouse chick were t r a n s p l a n t e d  s k i n of an 80-day o l d male D. o. p a l l i d u s .  under  110 In 1960, had  22 t h i r d - s t a g e larvae from a s i m u l i i d that  fed on grouse #25  (harboring Mf.  sp. B)  twenty-one days  e a r l i e r were i n j e c t e d i n t r a v e n o u s l y i n t o a 58-day o l d male D.  o.. p a l l i d u s c h i c k hatched i n c a p t i v i t y . In subsequent years  no  further transmission  attempts  were c a r r i e d out, e i t h e r because of the s c a r c i t y of inoculum or because young c h i c k s were not a v a i l a b l e . 8. C o l l e c t i o n of Data on N a t u r a l At the beginning to maintain serial  Transmission  of t h i s  study  the w r i t e r planned  u n i n f e c t e d grouse, some c o n t i n u o u s l y ,  i n t e r v a l s , throughout the s p r i n g and  summer on  grouse range, where they would be v u l n e r a b l e n a t u r a l l y i n f e c t e d vectors of Sk. and Mf.  sp. B.  provide  data  I t was  i n the  throughout the microfilariae  s p r i n g and  aetiology.  This plan -could  and  and  examined f o r  worms i n order to determine  at which they had  indoors  was  Grouse c h i c k s were c o l l e c t e d  summer of 1959-1961 and  and developing  c h i c k s were shot viscera  f l e x i v a g i n a l i s , S. p e c t o r a l i s  become i n f e c t e d .  a l i v e , most of them l e s s than two  maintained  by  p e r t a i n i n g to n a t u r a l t r a n s m i s s i o n  f o l l o w i n g ways.  approximate dates captured  the  out.  Information obtained  to attack  at  hoped that grouse so exposed would  on e p i z o o t i o l o g y and  not be c a r r i e d  others  field,  Those  weeks o l d , were  tested, p e r i o d i c a l l y u n t i l  examined i n the  the  and  killed. smears  and  from j u v e n i l e s shot by hunters c o n t r i b u t e d f u r t h e r  information.  Older  Ill  RESULTS AND DISCUSSION I.  A. Incidence  of F i l a r i a s e s i n G a l l i n a c e o u s  In t h i s in  DISTRIBUTION AND PREVALENCE OF NATURAL FILARIAL INFECTIONS  s e c t i o n the occurrence  the G a l l i f o r m e s of B r i t i s h Columbia  presented, regions.  of f i l a r i a l i n f e c t i o n s  and Alaska w i l l be  based on examinations of these  hosts  d e t a i l here,  Such species w i l l not be t r e a t e d - i n  and the reader  i s r e f e r r e d to Appendix-Tables  I I , and I I I , f o r a summary of a l l f i l a r i a l It  filarioids  from s e v e r a l  Few i n d i v i d u a l s of some species of g a l l i n a c e o u s  b i r d s were examined.  I,  Birds  was pointed  i n f e c t i o n s found.  out i n Part One that two species of  i n the g a l l i n a c e o u s b i r d s of B r i t i s h Columbia  duce m i c r o f i l a r i a e which are so s i m i l a r that they d i f f e r e n t i a t e d e f f e c t i v e l y on morphological be  cannot be  grounds.  I t could  argued, i n c o n t e n t i o n with t h i s i n t e r p r e t a t i o n , that  m i c r o f i l a r i a e represent  that these  these  only one s p e c i e s , the a d u l t s of which  occupy more than one s i t e . of M i c r o f i l a r i a  pro-  While, i n the absence of adults  sp. B, i t cannot be proved beyond a doubt  microfilariae  are produced by two species  than one, the f o l l o w i n g i n f o r m a t i o n i s presented t h i s i n t e r p r e t a t i o n seems the more p r e f e r a b l e :  rather  to show why l ) On Vancouver  I s l a n d , S p l e n d i d o f i l a r i a p e c t o r a l i s a d u l t s were never found in  d e t a i l e d examination of 37 grouse which harbored s h a r p - t a i l e d  * As used here, "prevalence" frequency of i n f e c t i o n s i n a host species from a given r e g i o n ; " i n c i d e n c e " = prevalence and d i s t r i b u t i o n i n v a r i o u s hosts and r e g i o n s . =  112 m i c r o f i l a r i a e , nor i n s u p e r f i c i a l ' examination of hundreds of grouse skinned by other workers. adults  2) In the 34 grouse (19  and y e a r l i n g s ; l b j u v e n i l e s ) from c e n t r a l B r i t i s h  Columbia which harbored s h a r p - t a i l e d m i c r o f i l a r i a e and were c a r e f u l l y searched f o r f i l a r i o i d s , 25-, and i t occurred  S. p e c t o r a l i s was/found i n  i n one s i t e only - under the breast  skin.  These f i n d i n g s i n d i c a t e s t r o n g l y that S-. p e c t o r a l i s does not occur on Vancouver I s l a n d , and, concomitantly, Mf. sp. B must be considered  that  distinct.  1. Occurrence on Vancouver I s l a n d a. Prevalence of S k r i abinocta The filariae other  w r i t e r has found no i n d i c a t i o n that the micro-  of t h i s  species  flexivaginalis  species  can be confused with those' of any  i n Galliformes  of t h i s r e g i o n ;  therefore,  diag-  n o s i s of i n f e c t i o n s with Sk. f l e x i v a g i n a l i s i s based on microfilaraemias  as w e l l as on presence of a d u l t s .  On Vancouver I s l a n d the w r i t e r found Sk. f l e x i v a g i n a l i s i n f e c t i o n s prevalent f i l a r i o i d was present Queen C h a r l o t t e  i n adult blue  grouse  (see Table I X ) .  This  i n the 2 adult D. o. s i t k e n s i s from the  Islands  and was .sporadic  i n ( l o f 9 adult)  r u f f e d grouse from south c o a s t a l B. C. Almost a l l the grouse i n which Mf. f l e x i v a g i n a l i s was found, and which were searched c a r e f u l l y (about 4 0 ) , y i e l d e d adults of t h i s occupied  species.  The considerable  d i v e r s i t y of s i t e s  by these worms worms renders i t u n l i k e l y that a l l  a d u l t s were recovered  from each i n f e c t e d grouse. /The maximum  number found was 10 males and 14 females; but the average'  113 infection  was l i g h t , 2 males and 3 females.  always r a t h e r t i g h t l y  coiled  i n small thin-walled, cysts  a s s o c i a t e d with the w a l l of a blood v e s s e l . i n the f o l l o w i n g s i t e s ,  The worms were usually  Adults were found  l i s t e d i n order of frequency of occu-  pation. 1)  Primary t r i f u r c a t i o n  of s u p e r i o r mesenteric  2)  Beside c o e l i a c  3)  Beside s u p e r i o r mesenteric  4)  Beside d o r s a l a o r t a , near c o e l i a c lungs ( 4 ) .  5)  W i t h i n the adrenal glands- u n i l a t e r a l  6)  Beside i s c h i a t i c v e i n  7)  Beside branches  8)  Beside i n f e r i o r mesenteric  9)  Beside i n t e s t i n a l v e i n ( l ) .  a r t e r y near d o r s a l aorta ( 4 ) . a r t e r y , at base of  -  (2); b i l a t e r a l ( l ) .  (2).  of s u p e r i o r mesenteric  artery  (2).  artery ( l ) .  D o r s a l to middle  11)  Between femoral a r t e r y , v e i n , and nerve ( l ) .  12)  Undetermined ( 3 ) .  of vas deferens ( l ) .  b. Prevalence of M i c r o f i l a r i a On Vancouver I s l a n d t h i s  sp. B s p e c i e s i s very p r e v a l e n t  (see Table IX) but i t occurred i n only 4 of 9  adult r u f f e d grouse sula) .  (7) .  artery (6).  10)  i n blue grouse  artery  ( i n c l u d i n g one b i r d  from the Sechelt Penin-  Mf. sp. B was present i n 1 of the 2 adult blue  grouse  from the Queen C h a r l o t t e I s l a n d s . c. Comparative prevalence of these 2 s p e c i e s on Vancouver I s l a n d It can be seen from Appendix Table I that 96% of the adult blue grouse  examined from Vancouver I s l a n d had  114 microfilariae  i n t h e i r blood; i n 67% of these i n f e c t i o n s both  Mf.  f l e x i v a g i n a l i s and Mf. sp. B occurred.  the  differences i n habits  and h a b i t a t  In the l i g h t of  s e l e c t i o n that e x i s t  between males and females and between a d u l t s worth c o n s i d e r i n g incidence lence  i t is  whether r e l a t e d d i f f e r e n c e s i n m i c r o f i l a r i a l  also e x i s t .  There was l i t t l e  among the 3 regions  from the 3 regions  d i f f e r e n c e i n preva-  from which grouse were c o l l e c t e d :  Nanaimo Lakes, Courtenay, and Quinsam.  picture.  and chicks  In Table IX the data  have been pooled t o provide a comprehensive  I t should be noted that because the y e a r l i n g sample  i s very s m a l l , 5 y e a r l i n g s which had spent 2 summers on the grouse range have been i n c l u d e d only  with the 11 which had spent  one summer. Table IX.  M i c r o f i l a r i a l i n f e c t i o n s i n the blue grouse examined from Vancouver I s l a n d (56 adult males, 29 adult females, and 16 y e a r l i n g s ) .  Males No. Inf.  Females % Inf.  No. Inf.  Yearlings  % Inf.  No. Inf.  % Inf.  Microfilariae  56  100  26  90  12  75  Both Mf. sp. B and Mf. f l e x .  40  71  15  52  4  25  Mf.  sp. B only  10  18  6  21  6  38  Mf.  flex,  11  5  17  2  12  Mf-  sp. B  only  Mf. f l e x i v a g i n a l i s  6 50  89  21  72 ' ,  46  82  20  69  * Eight males and e i g h t females pooled because there d i f f e r e n c e i n i n f e c t i o n s between sexes.  10 6  was no  62 38  115 The noteworthy f e a t u r e s be summarized as f o l l o w s :  l ) I n f e c t i o n s with Mf. sp. B are  s i g n i f i c a n t l y more p r e v a l e n t (z=1.67) or i n y e a r l i n g s d i f f e r e n t between adult  of the data i n Table IX can  i n adult males than i n adult  females  (z=6.4), but are not s i g n i f i c a n t l y females and y e a r l i n g s  (z=.35).  2)  I n f e c t i o n s with Mf. f l e x i v a q i n a l i s are not s i g n i f i c a n t l y more prevalent  i n adult males than i n adult  s i g n i f i c a n t l y more p r e v a l e n t adult  females  ( z - l . l ) but are  i n adult males (z=3.2) and i n  (z=1.73) than i n y e a r l i n g s .  any sex or s t a t u s significantly  females  the two species  3) In grouse of  of f i l a r i o i d s do not d i f f e r  i n the prevalence of t h e i r i n f e c t i o n s .  These d i f f e r e n c e s i n prevalence are i n t e r p r e t e d to i n d i c a t e , from the standpoint  of the host, that  as c h i c k s , become i n f e c t e d with f i l a r i o i d s but and  they continue to acquire 2) a f t e r t h e i r f i r s t  contract  l ) most grouse,  in their first  i n f e c t i o n s i n subsequent  summer,  years,  year, males are more l i k e l y to  Mf. sp. B i n f e c t i o n s than are females: and from the  standpoint  of the p a r a s i t e s , that Mf. sp. B i s not  more s u c c e s s f u l l y than Sk.  transmitted  flexivaqinalis.  2. Occurrence of i n f e c t i o n s i n c e n t r a l and northern B r i t i s h Columbia and Alaska Appendix Table I I i n d i c a t e s that 55% of the 58 and  yearling tetraonids  examined from the i n t e r i o r of B. C.  harbored m i c r o f i l a r i a e , and i n 14% of these grouse the  i n f e c t i o n s , two o b v i o u s l y  present.  adult  (or 25%) of  d i f f e r e n t m i c r o f i l a r i a e were  Most of the i n f e c t i o n s were i n blue grouse, where  of the 35 adults of these b i r d s microfilariae  69%  and y e a r l i n g s harbored m i c r o f i l a r i a e and i n 18%  (25% of the i n f e c t i o n s ) two o b v i o u s l y occurred.  different  116 a. Incidence of S k r i abinocta  flexivaqinalis  In the i n t e r i o r of the province found i n 6 of 32 adult blue grouse  (adults otherwise),  grouse, i n 1 ( j u v e n i l e ) of 11 r u f f e d i n 2 of 9 adult F r a n k l i n ' s and i n  none of a small number of adult  and j u v e n i l e s h a r p - t a i l e d  grouse, chukar p a r t r i d g e , gray p a r t r i d g e b. Incidence of M i c r o f i l a r i a The 58  adult  t h i s f i l a r i o i d was  and C a l i f o r n i a  quail.  sp. B  data i n Appendix Table I I show that 34 (59ft) of  and y e a r l i n g grouse of 4 species  from non-coastal  harbored s h a r p - t a i l e d m i c r o f i l a r i a e .  E i g h t of these b i r d s  yielded  to y i e l d  adult S. p e c t o r a l i s . 9 f a i l e d  oping S. p e c t o r a l i s . and 17 were not autopsied were t e s t e d at Game-Checking S t a t i o n s .  adult or d e v e l because they  In some of these 17  grouse not searched, the p e c t o r a l y e l l o w i n g ,  c h a r a c t e r i s t i c of  heavy i n f e c t i o n s with S. p e c t o r a l i s . was present. only 8 of the 17 m i c r o f i l a r a e m i c opsied  B. C.  The f a c t  that  adult and y e a r l i n g grouse aut-  a c t u a l l y harbored adult S. p e c t o r a l i s suggests that the  m i c r o f i l a r i a e i n the remainder and i n some of those not searched may have been Mf. sp. B (but see pp. 128-129). c. Incidence of S p l e n d i d o f i l a r i a p e c t o r a l i s In the i n t e r i o r of B r i t i s h Columbia, S. p e c t o r a l i s i s moderately abundant i n D. o. p a l l i d u s i n the Trout  Creek  V a l l e y where i t was found i n 14 of 23 a d u l t s and j u v e n i l e s , and near Oyama - i n 7 of 17 a d u l t s , y e a r l i n g s , and j u v e n i l e s . 1 of 12 blue  grouse from 7 other  i n t e r i o r y i e l d e d these worms. Based on adult  localities  Only  i n the c e n t r a l  S. p e c t o r a l i s was a l s o found i n  and l a r v a l worms, not on m i c r o f i l a r i a e .  117 s h a r p - t a i l e d grouse  and spruce grouse  i n Alaska, but was not encountered i a n i d s examined. found  i n s o u t h - c e n t r a l B, C. and  i n the small number of phas-  I t i s noteworthy that no m i c r o f i l a r i a e were  i n 10 a d u l t r u f f e d grouse  from c e n t r a l B. C.  S. p e c t o r a l -  i s seems to be abundant i n Alask'a (see Appendix Table I I I ) , and it  i s probable  that i t s d i s t r i b u t i o n i s more or l e s s  continuous  from s o u t h - c e n t r a l B r i t i s h Columbia north to c e n t r a l Alaska. The  i n t e n s i t y of i n f e c t i o n s with S. p e c t o r a l i s i s  r a t h e r v a r i a b l e , sometimes being as high as 50 worms and not i n f r e q u e n t l y , s e v e r a l dozen. g e n e r a l s i t e - subcutaneously  They are c o n s i s t e n t l y i n one  i n somewhat t a n g l e d  aggregations  about midway between the crop and the xiphisternum, slightly  to the r i g h t of the m i d - v e n t r a l  d. Incidence of M i c r o f i l a r i a In North America,  this  line.  lagopodis  (?-S. p a p i l l o c e r c a )  species has been found  i n extreme northern B r i t i s h Columbia and In Alaska only i n ptarmigan. w i l l o w ptarmigan w r i t e r found and  In Alaska, Babero (8) found  infected.  ptarmigan  were n e g a t i v e .  f a r only Mf. lagopodis N e a r c t i c ptarmigan,  ptarmigan  yet 10 adult, w h i t e - t a i l e d occurred  from e a s t - c e n t r a l A l a s k a .  I t i s i n t e r e s t i n g to note  that so  (? S. p a p i l l o c e r c a ) has been found i n  and that i t has not been found i n other  Nearctic tetraonids. this f i l a r i o i d  B. C. the  However, adult S. p a p i l l o c e r c a  i n one mature w h i t e - t a i l e d ptarmigan Interpretation.  ( 8 ) , and  i n 8 of 20 adult w i l l o w  i n 1 of 17 adult rock ptarmigan,  only  19 of 42  In extreme northwestern  this microfilaria  usually  This seems to imply that i n North  America  i s r e s t r i c t e d to the genus Lagopus and that the  3 other f i l a r i o i d s do not p a r a s i t i z e ptarmigan,  even though  118  they do occur i n t e t r a o n i d s of 3 other tions require  that Mf.  (see p. 44). lagopodis  a b a s i c premise:  as reported  we  that  The  Mf.  w r i t e r suggested e a r l i e r  (p.  I f we  i n the  northern  of Splendido^  l) i n western Europe lagopodis  2 H o l a r c t i c species  Nearctic  occurs ,in s e v e r a l i n North America  of Lagopus: and  ( i n c l u d i n g Lagopus) and  only  i n the  2)  that i n  one  Since  but  specificity  i s no reason apparent why  i t should  I f lagopodis  i s not  i n Asia  p a p i l l o c e r c a then  L. leucurus  from other  Thus the  The  Let us For c l a r i t y  this  the leucurus writer implies  or a l t i t u d i n a l i s o l a t i o n  ptarmigan than a c t u a l l y e x i s t s  available information  hypothesis that Mf.  cerca.  an e c o l o g i c a l s p e c i f i c i t y .  degree of g e o g r a p h i c a l  laoopodis  there  behave d i f f e r e n t l y i n  that such a r e s t r i c t e d e c o l o g i c a l s p e c i f i c i t y  a greater  species  (see p. 34)  r e s t r i c t i o n of p a p i l l o c e r c a . i n North America, to L. presumably represents  i n the  S. p a p i l l o c e r c a shows a  broad p h y s i o l o g i c a l host  North America.  phasianid,  sole r e s t r i c t e d l y North American  of Lagopus (L. l e u c u r u s ) .  p. 99).  true,  A s i a p a p i l l o c e r c a occurs i n s e v e r a l genera of  tetraonids  thinks  46)  hypothesize that t h i s i s not  genera of t e t r a o n i d s i n c l u d i n g Lagopus. and only  the  elsewhere as  i s probably the m i c r o f i l a r i a  papillocerca.  that  i n ptarmigan i n northwestern North  same species  filaria find  set out  found to-date  America i s the lagopodis  These i m p l i c a -  c l o s e r examination.  Let us f i r s t microfilaria  genera.  fails  of  (see  to support  an  i s not S. p a p i l l o c e r c a .  assume, t e n t a t i v e l y , that they are one species w i l l be r e f e r r e d to here as S.  Working from t h i s assumption we  see  species. papillo-  that S. p a p i l l o c e r c a  119 i s a H o l a r c t i c species which p a r a s i t i z e s a number of  galliform  genera i n the P a l e a r c t i c , but p a r a s i t i z e s only the 3 species of Laqopus i n the N e a r c t i c . r e s t r i c t i o n of t h i s f i l a r i o i d attributable  to one  specificity.  Since  In North America the  to ptarmigan i s l i k e l y  of the f o l l o w i n g f a c t o r s :  l) E c o l o g i c a l  the summer range of ptarmigan i s u s u a l l y  somewhat above that of other te.traonids there may opportunity  apparent  be very  f o r t r a n s m i s s i o n of S. p a p i l l o c e r c a to these  little other  species.  2) P h y s i o l o g i c a l s p e c i f i c i t y .  incapable  of i n f e c t i n g N e a r c t i c t e t r a o n i d s other than Laqopus  its  I f S. p a p i l l o c e r c a i s  a b i l i t y to i n f e c t P a l e a r c t i c t e t r a o n i d s other than Laqopus  might imply  that the P a l e a r c t i c t e t r a o n i d s are more c l o s e l y  r e l a t e d to each other  and  to the- species of Laqopus i n  northern  A s i a than are the N e a r c t i c t e t r a o n i d s to species of Laqopus i n North America.  3) I n s u f f i c i e n t  sampling.  t h a t an e f f o r t be made to examine L. l e u c u r u s . and Alaska  and  overlap.  w r i t e r suggests  a) ptarmigan, p a r t i c u l a r l y  b) other t e t r a o n i d s , i n those  northern  regions  of  B r i t i s h Columbia where t h e i r summer ranges  By so doing,  i t should be p o s s i b l e to determine  g r e a t e r c e r t a i n t y whether the (Mf.  The  with  "ptarmigan m i c r o f i l a r i a "  l a q o p o d i s ) * i s that of S. p a p i l l o c e r c a . and  whether  other t e t r a o n i d s harbor S. p a p i l l o c e r c a when c o n d i t i o n s  favour  t h e i r being b i t t e n by i t s i n f e c t e d v e c t o r s .  Since the a d u l t s of Mf_. laqopodis were not found by Brinkmann ( l o c . c i t . , Part One), who d e s c r i b e d t h i s m i c r o f i l a r i a the most completely, i t w i l l probably never be proved c o n c l u s i v e l y that h i s m i c r o f i l a r i a was Mf. p a p i l l o c e r c a . If t h i s could be done, the name laqopodis would take p r i o r i t y over p a p i l l o c e r c a .  120 B.  Incidence i n Non-Gallinaceous  Birds  Appendix Table IV presents the r e s u l t s of sampling the n o n - g a l l i n a c e o u s avifauna on and near grouse range. t o t a l of 225  adult b i r d s was  i n 27 f a m i l i e s .  v i n c e , to 11.4%  examined, r e p r e s e n t i n g 79 s p e c i e s  M i c r o f i l a r i a l i n c i d e n c e ranged  i n d i v i d u a l s and 5.9%  A  from 3.7%  of the  of the s p e c i e s i n the i n t e r i o r of the pro-  of the i n d i v i d u a l s and 22% of the s p e c i e s from  Vancouver I s l a n d .  Appendix Table V p r o v i d e s b r i e f  quantitative  d e s c r i p t i o n s and Appendix VI' summarizes the q u a l i t a t i v e f e a t u r e s of the m i c r o f i l a r i a l  forms encountered.  Most of these are sep-  arable from those of the t e t r a o n i d s by a combination of characters. Mf.  However, two  flexivaginalis:  forms from Vancouver I s l a n d c l o s e l y Form 3 - a long, b l u n t - t a i l e d  microfilaria  from a M a c G i l l i v r a y ' s warbler; and Form 4 - a s h o r t e r aria  from Audubon's warbler and towhee.  filariae  are compared with Mf.  resemble  microfil-  In Table X these micro-  flexivaginalis.  The p o s i t i o n s of  f i x e d p o i n t s are s t r i k i n g l y s i m i l a r ; however, Form 3 can be differentiated  from Mf.  f l e x i v a g i n a l i s by g r e a t e r l e n g t h , a  much longer sheath, and by the c l e f t  appearance  and s u b a p i c a l  p o s i t i o n of the p o s t e r i o r end of the n u c l e a r column. may  be one of the s p e c i e s i n o l i v e - s i d e d  grosbeak,  Form 4  f l y c a t c h e r and  but m i c r o f i l a r i a e of t h i s type i n smears.from  hosts were scarce and not w e l l s t a i n e d .  evening these  The w r i t e r can f i n d  means of d i s t i n g u i s h i n g between Form 4 and Mf.  no  flexivaginalis.  but because a d u l t s of Form 4 were not found i n l o c a l -xpasserines t h i s m i c r o f i l a r i a cannot be p o s i t i v e l y i d e n t i f i e d .  It should be r e c a l l e d that S k r i abinocta f l e x i v a g i n a l i s d e s c r i b e d from crows i n Ohio (loci'; c i t . . Part One).  was  Table X.  Measurements* of s i m i l a r microf i l a r i a e from grouse and p a s s e r i n e s .  Mf. f l e x i v a q i n . Host  Grouse  No. of Mf.  85  Form 3. 0.  tolmiei 10  Form 4 D. auduboni  Form 4 Pioilo  20  10  170 ( 139-228 )  276 ( 235-342 )  205 ( 186-216 )  165 ( 150-170 )  To NR 1{%)  24.4(21.6-26.9)  24.7(23 .6-26.0)  24.9(23.1-27.6)  23.7(21.4-26.9)  To EV 1[%)  36.0(33.6-40.0)  35.2(34.1-36.3)  36.3(35.2-37.2)  35.8(34.2-38.9)  To I B  {%)  63.4(58.8-64.4)  64.0(61.2-66.1)  63.8(61.8-67.5)  63.5(61.5-66.2)  {%)  67.4(61.5-72.2)  66.8(64.4-68.2)  67.0(65.3-71.3)  66.7(63.8-69.3)  '[%)  76.3(72.3-81.4)  76.3(74.9-78.0)  77.6(74.8-8.1.2)  78.7(76.4-80.6)  To AV 1:%)  89.5(86.5-91.4)  89.8(85.6-92.8)  89.3(88.5-90.6)  89.7(87.8-91.4)  Length  To I B To R  x  (u.)  L  2  * D i s t a n c e s to f i x e d p o i n t s expressed as percentages of body l e n g t h .  122 C.  Summary and T e n t a t i v e The  prevalence  b i r d s varies*markedly w i t h i n a host host  Proposals of f i l a r i a l  infections i n gallinaceous  among hosts of d i f f e r e n t  species p o p u l a t i o n , among p o p u l a t i o n s  regions.  d i s t r i b u t i o n deserve  The f o l l o w i n g aspects  of prevalence and  further attention:  1) The absence of S. p e c t o r a l i s from south 2) The absence of f i l a r i o i d s  of grouse  c o a s t a l B. C.  (except p o s s i b l y  f l e x i v a q i n a l i s ) i n n o n - t e t r a o n i d b i r d s of the same r e g i o n . 3) The d i f f e r e n t  of  of the same  s p e c i e s , among the s p e c i e s of hosts, and between  geographic  Sk.  sex and age  filarial  prevalences  w i t h i n the Tetraonidae.  4)  The low prevalence  5)  In Vancouver I s l a n d blue grouse, the g r e a t e r  filariases  of Sk. f l e x i v a q i n a l i s i n c e n t r a l B. C. prevalence  of Mf.- sp. B i n a d u l t males than i n adult females  and y e a r l i n g s , as opposed to the g r e a t e r prevalence v a q i n a l i s i n f e c t i o n s i n a d u l t s than In the f i n a l  chapter  of Sk. f l e x i -  in yearlings.  of the t h e s i s an attempt w i l l be  made to account f o r the d i f f e r e n c e s i n i n c i d e n c e which have been shown to e x i s t .  At present  i t can be seen that a c o n s i d e r a b l e  number of f a c t o r s must i n f l u e n c e the d i s t r i b u t i o n and prevalence of  the f i l a r i o i d s  found  a "vector i s necessary of  these  i n B r i t i s h Columbia t e t r a o n i d s .  i n the l i f e  f a c t o r s are probably  c y c l e s of f i l a r i a l  related  Since  worms, most  to t r a n s m i s s i o n .  In the f o l l o w i n g s e c t i o n the w r i t e r w i l l  t r y to-esta-  b l i s h when t r a n s m i s s i o n takes p l a c e , as i n d i c a t e d by the occurrence  of young i n f e c t i o n s , and s e v e r a l aspects  of the f i l a r i a s e s  themselves which i n f l u e n c e t h e i r t r a n s m i s s i o n w i l l  be c o n s i d e r e d .  123  II.  A.  OTHER INFORMATION DERIVED FROM NATURAL FILARIAL INFECTIONS  I n d i c a t i o n s of Transmission Period. 1.  On Vancouver I s l a n d Bendell  blue grouse  (10)  failed  to d e t e c t m i c r o f i l a r i a e i n  157  c h i c k s shot i n June, J u l y and August, 1950-54, i n  the Quinsam r e g i o n .  Between June 11 and J u l y 1, 1958-61, the  w r i t e r , with the a s s i s t a n c e of o t h e r s , captured about 90 blue grouse  c h i c k s from  c h i c k s ranged  the Quinsam and Nanaimo Lakes areas.  i n age  from 2 days to 3 weeks when captured,  the p e r i p h e r a l blood of a l l of them was ariae.  These and  negative f o r m i c r o f i l -  They were subsequently maintained  c o n d i t i o n s , and when they were s a c r i f i c e d  under f l y - p r o o f or d i e d , u s u a l l y  w i t h i n a year of capture, post mortem examination demonstrated that m i c r o f i l a r a e m i a s were absent  of lung blood  from a l l of these  birds. Blue grouse and  1961  c h i c k s shot during the summers of  at Nanaimo Lakes,  i n c l u d i n g s i x 80-day o l d and  1960 two  55-day o l d b i r d s c o l l e c t e d August 29-31, I960, were a l s o negative f o r m i c r o f i l a r i a e grouse  chick,  J u l y 27, vity  1961  and developing worms.  One  blue  about 50 days o l d when captured at Quinsam, was  a l s o negative throughout  and on autopsy  9 months a f t e r  i t s p e r i o d of c a p t i -  capture.  In the l i g h t of t h i s overwhelming evidence  that mi-  c r o f i l a r a e m i a s do not become patent i n Vancouver I s l a n d grouse c h i c k s before summer's end, Mf.  flexivaginalis  and Mf.  i t was  s u r p r i s i n g to f i n d  sp. B i n a smear from  both  a 58-day o l d  124 c h i c k shot August 21, Bendell). grouse  1959  F u r t h e r evidence  in their first  microfilariae  of SJ<.  Mf.  flexivaqinalis  two  collected  ( c o l l e c t i o n of Dr. J . F.  of t r a n s m i s s i o n of f i l a r i o i d s  summer was  the presence  1960  at Nanaimo,  and/or Mf.  sp.  ;  l ) atiults  and  grouse  B i n 9-month o l d j u v e n i l e s , collected April  the other taken A p r i l 5, 1960  There i s i n d i r e c t  to  2) m i c r o f i l a r a e m i a s of  from Quinsam i n March 1960, one  at Courtenay,  of  f l e x i v a q i n a l i s i n a j u v e n i l e blue  c o l l e c t e d November 30,  1960  at Quirisam  evidence  at Naaaimo  that t r a n s m i s s i o n of  17, Lakes.  Mf.  sp. B can occur, though i n f r e q u e n t l y , before the end of June. •i  l ) Bendell (unpublished data) i n June harbored  Leucocvtozoon  found that 2 of 41 c h i c k s examined sp;'  2) Subsequently,  Woo  (55)  demonstrated t h a t sporogony of a s p e c i e s of Leucocytozoon grouse  from  can occur i n g r o u s e - b i t i n g s i m u l i i d s .  2.  In the Okanagan  Region  In the s o u t h - c e n t r a l part of the p r o v i n c e , where S. p e c t o r a l i s o c c u r s , i n d i c a t i o n s of" the p e r i o d of n a t u r a l transmiss i o n of t h i s  s p e c i e s were r e a d i l y obtained because these worms  occupy a r e s t r i c t e d , e a s i l y examined S i x t e e n blue grouse  site.  and 6 r u f f e d grouse  c h i c k s , 2 days  to 4 weeks o l d , were captured at Trout Creek and Oyama between June 2 and June 18, filaraemias ral  s k i n and  autopsy. and  failed  two,  1959,  and maintained  to develop  in captivity.  Micro-  i n any of these b i r d s and the  abdominal blood v e s s e l s of a l l were negative  Three other blue grouse' c h i c k s shot during June about 33 days o l d , shot J u l y 2, 1959 R e s u l t s of examination  l a t e r i n the summers of 1958,  and  1960  on 1959  were a l s o n e g a t i v e .  of blue grouse  1959,  pecto-  chicks c o l l e c t e d  from 2 l o c a t i o n s i n  125  the i n t e r i o r of the province were as f o l l o w s : At Trout Creek - a l l but one were blue or  1) J u l y 21, 1959 - 50-day o l d female. microfilariae.  or  2) J u l y 24, 1960 - 50-day o l d male. microfilariae.  grouse.  No developing worms No developing worms  3) J u l y 25, 1959 - 50-day o l d male. Beneath p e c t o r a l s k i n were 11 f o u r t h - s t a g e l a r v a e , 1 female (1/3 adult l e n g t h ) , and 2 f u l l y grown males (no sperm) - a l l S. p e c t o r a l i s . No microf i l a r i a e i n blood. 4) J u l y 25, 1959 - 50-day o l d female (same brood). p e c t o r a l s k i n were 10 f o u r t h - s t a g e larvae and 1 female length) - S. p e c t o r a l i s . No m i c r o f i l a r i a e i n blood. 5) August 1, 1959 - two 30-day o l d females No developing worms or m i c r o f i l a r i a e .  Beneath (1/2 adult  (from one brood).  6) August 3, 1960 - 60-day o l d male. Beneath p e c t o r a l was a p a i r of a d u l t - s i z e d , but immature S. p e c t o r a l i s . No m i c r o f i l a r i a e i n blood.  skin  7) August 11, 1959 - 75-day o l d male. Beneath p e c t o r a l s k i n were 25 a d u l t - s i z e d S. p e c t o r a l i s . ( T r a n s f e r experiment). No m i c r o f i l a r i a e i n blood. 8) August 11, 1959 - 75-day o l d male (same brood). Maint a i n e d i n c a p t i v i t y u n t i l February 8, 1960. P e r i p h e r a l blood negative throughout c a p t i v i t y . At necropsy, lung blood with s h a r p - t a i l e d m i c r o f i l a r i a e ; p e c t o r a l s k i n with 2 encapsulated dead females of S. p e c t o r a l i s . 9) August 19, 1959 - 85-day o l d male. Beneath p e c t o r a l s k i n were 31 a d u l t - s i z e d S. p e c t o r a l i s ; some males producing sperm. No m i c r o f i l a r i a e i n females or i n blood. 10) August 19, 1959 - 85-day o l d female (same brood). Beneath p e c t o r a l s k i n were 46 a d u l t - s i z e d .S. p e c t o r a l i s ; some males producing sperm. No m i c r o f i l a r i a e i n females or i n blood. 11) September 1, 1959 - ca. 90-day o l d male. Beneath p e c t o r a l s k i n were 23 male and 25 female adult S. p e c t o r a l i s . M i c r o f i l a r i a e in. utero and i d e n t i c a l m i c r o f i l a r i a e i n blood. 12) September 1, 1959 - ca. (?)80-day o l d male r u f f e d grouse. Beneath p e c t o r a l s k i n were 1 male and 2 female adult S. pecto r a l i s . M i c r o f i l a r i a e i n utero and i d e n t i c a l m i c r o f i l a r i a e i n blood.  126 At Oyama (12 miles south of Vernon, between Okanagan and Kalamalka Lakes, e l e v a t i o n 3000 feet) - a l l were blue grouse. or  1) J u l y 30, 1958 - 11-week o l d male. microfilariae.  No developing worms  2) J u l y 30, 1958 - 9-week o l d male. Beneath p e c t o r a l s k i n were 2 male and 6 female a d u l t - s i z e d S . p e c t o r a l i s . ' No microf i l a r i a e i n u t e r i or blood. 3) J u l y 30, 1958 - 9-week o l d female. Beneath p e c t o r a l s k i n were 3 a d u l t - s i z e d male S. p e c t o r a l i s . No m i c r o f i l a r i a e . 4) J u l y 30, 1958 - 9-week o l d female. Beneath p e c t o r a l s k i n were 7 male and 5 female a d u l t - s i z e d S. p e c t o r a l i s . Males producing sperm; m i c r o f i l a r i o i d embryos present, but not f u l l y developed, i n 1 female. No m i c r o f i l a r i a e i n blood. or  5) August 26, 1958 - 14-week o l d male. microfilariae.  No developing worms  6) August 27, 1958 - 11-week o l d female. Beneath p e c t o r a l s k i n were 1 male and 1 female adult S. p e c t o r a l i s . Microfilariae i n u t e r o . blood n e g a t i v e . 7) August 27, 1958 - 12-week o l d male. Beneath p e c t o r a l s k i n were 1 male and 1 female adult S. p e c t o r a l i s . . M i c r o f i l a r i a e i n u t e r o , blood negative. Four blue grouse hens were c o l l e c t e d time as four of the chicks hens) that  yielded  to note that  at the same  (apparently the o f f s p r i n g  developing S. p e c t o r a l i s .  of these  It i s interesting  none of these hens harbored adults or developing  larvae of S. p e c t o r a l i s . Sharp-tailed  or any m i c r o f i l a r i a e . microfilariae  of the " p e c t o r a l i s  type were found i n blood smears of 7 of the 21 j u v e n i l e ruffed,  and s h a r p - t a i l e d  i n September of s e v e r a l to d i s t i n g u i s h  grouse tested years.  Since  blue,  at Game-Checking S t a t i o n s 1) the w r i t e r  between Mf. sp. B and Mf. p e c t o r a l i s .  was not p o s s i b l e  - sp. B"  to examine the subcutaneous p e c t o r a l  i s unable and  2)  it  t i s s u e s of  127 these b i r d s , there was  no way  of determining  both) species had been t r a n s m i t t e d to these  which (or whether birds..  Of a l l the j u v e n i l e grouse.examined from the c e n t r a l i n t e r i o r of the province only one, the southern. Cariboo, September 20, Sk.  f l e x i v a g i n a l i s adults.  a r u f f e d grouse from was  found  to  harbor  In view of the s c a r c i t y of t h i s  s p e c i e s i n the i n t e r i o r of B. C., and  1959,  south-  and p a r t i c u l a r l y  i n the f a c t that the male and 2 females  were  i n Bonasa.  accidentally  d i s l o d g e d from f o r m a l i n - p r e s e r v e d v i s c e r a , t h e i r d i s c o v e r y rather fortuitous. "pectoralis-sp.  Lung blood from t h i s b i r d contained  B" m i c r o f i l a r i a e , but  oped m i c r o f i l a r i a e were present B.  I n d i c a t i o n s of Prepatent  was  only  apparently f u l l y d e v e l -  i n the u t e r i of the 2  females.  Period  This i n t e r v a l , the length of time  e l a p s i n g between  i n o c u l a t i o n of t h i r d - s t a g e larvae and d e t e c t i o n of m i c r o f i l a r i a e i n the p e r i p h e r a l blood, i s r a t h e r v a r i a b l e among the s p e c i e s of f i l a r i o i d s which have been s t u d i e d . the minimum prepatent Litomosoides  carinii  80 days i n B. malayi and D. repens  For f i l a r i a e  of mammals  periods range from about 50 days i n (51), 59 days i n Brugia pahangi  (50), and  (22), to 6 months i n D i r o f i l a r i a  immitis  (52), and  11 months i n D.  aethiops  m i c r o f i l a r a e m i a s of C o n i s p i c u l u m . f l a v e s c e n s  (31).  in lizards  The are  d e t e c t a b l e as e a r l y as 41 days (36), while the prepatent of the avian f i l a r i o i d s Anderson  periods  s t u d i e d are even s h o r t e r .  (4) observed  microfilariae  of S.  fallisensis  i n ducks i n f e c t e d e x p e r i m e n t a l l y 30 days b e f o r e , and H i b l e r determined  (52)  that the minimum prepatent  periods f o r E u f i l a r i a  (33)  128 lonqicaudata.  Skriabinocta  s t r i a t o s p i c u l a and S p l e n d i d o f i l a r i a  p i c a c a r d i n a were 34 to 62 days. Field the present pectoralis Mf.  observations  of young i n f e c t i o n s i n grouse i n  study suggest that the prepatent p e r i o d  f o r S.  i s about 6 weeks, and f o r Sk. f l e x i v a g i n a l i s and  sp. B about 2 months, c a l c u l a t e d from probable date of  hatching.  For none of the grouse chicks harboring m i c r o f i l a r i a e  was the p r e c i s e date of i n f e c t i o n known,' and i t i s p o s s i b l e that experimental i n f e c t i o n would show that the prepatent periods C.  are a c t u a l l y s h o r t e r  Longevity The  than i n d i c a t e d .  of I n f e c t i o n s w r i t e r ' s observations  with S. p e c t o r a l i s may not be very  suggest that i n f e c t i o n s long-lived.  The host  reaction  to the presence of S. p e c t o r a l i s a d u l t s i s r a t h e r i n t e n s e , i n i n f e c t i o n s of only a few weeks' d u r a t i o n , found many worms surrounded i n s e v e r a l places connective imately  tissue.  One j u v e n i l e blue  even  and the w r i t e r has by strands of  grouse which died  approx-  6 months a f t e r becoming i n f e c t e d y i e l d e d the p a r t i a l l y  decomposed, p a r t l y encapsulated remains of s e v e r a l adult S. p e c t o r a l i s - presumably the worms r e s p o n s i b l e level microfilaraemia  i n this bird.  grouse from B. C. and Alaska  f o r a very low  In s e v e r a l s h a r p - t a i l e d  extensive  involvement and d e s t r u c -  t i o n of adult S. p e c t o r a l i s by f i b r o u s connective noted.  These observations  suggest that i n some  t i s s u e was instances  complete d e s t r u c t i o n of these f i l a r i o i d s may occur before f i l a r a e m i a s have d e c l i n e d a p p r e c i a b l y .  micro-  This, i n turn, r a i s e s  the p o s s i b i l i t y that at l e a s t some of the grouse from c e n t r a l B. C ,  i n which the presence of s h a r p - t a i l e d m i c r o f i l a r i a e and  129 the  apparent lack  actually  of adult  suggested Mf.  showed orphaned m i c r o f i l a r a e m i a s of .S,  A d u l t s or f o u r t h - s t a g e juvenile  19  adult  indication  from l a t e J u l y  blue grouse.  that  sp.  of the  on,  but  pectoralis  are  32  occurred  This i s construed  adults of S.  B,  pectoralis.  larvae were present i n 16  blue grouse c o l l e c t e d  i n only 5 of further  filarioids  as  rather  short-lived. In Sk. found no  f l e x i v a q i n a l i s i n f e c t i o n s , the  indication  todes, nor  of t i s s u e  of dead worms, and  duals of t h i s species are D.  Microfilarial  p e r i o d March 2-3, 1961.  63  was  and  It was  between 7 and  rather  tested  per  20  between any  0700 to  1900  microfilaria1  that  hours  low  every 3 hours f o r a 36-hour  counts of Mf.  exists  that,  sp." B ranged  rhythm.  natural  blood without The  greatest  12-hour averages of counts  (av.  to 0700 hours  count 45  each 24  mf.).  fluctuate  results  rhythm of Mf.  sp.  count 23  some i n d i c a t i o n peripheral  t h i s grouse B,  the  the  blood  may  possibility  during c a p t i v i t y , a c l a s s i c a l p e r i o d i c i t y  become subdued by  mf.)  between a c l e a r l y  hours there was  of t e s t i n g  (av.  was  Thus, although  were more numerous i n the  While the  represent the  two  periods 1900  microfilariae  diurnally.  indivi-  long-lived.  cu.mm. of p e r i p h e r a l  d e n s i t y d i d not  d e f i n e d high and  nema-  every 2 hours f o r a 24-hour p e r i o d March  found that  found between the and  the  i t seems probable that  showing a c l e a r l y marked c i r c a d i a n difference  involving  has  Periodicity  Grouse #24  6-7,  reaction  writer  prolonged i n a c t i v i t y of the  had  caged host.  130 During the second and t h i r d weeks of June, caged grouse used f o r exposures  1961,  the  near Nanaimo Lakes were housed  w i t h i n the r e s e a r c h c a b i n , where they were s u b j e c t e d u n i n t e n t i o n a l l y to a r t i f i c i a l each day. #24  A simuliid  (with Mf.  light  f o r 2 to 5 hours- a f t e r  sunset  (S. aureum) captured from,blue  sp. B) at 1730  grouse  hours PST on June 4 had i n g e s t e d  20 m i c r o f i l a r i a e , however two weeks l a t e r three S. aureum had only 2 m i c r o f i l a r i a e b i r d at 1700 liids  each i n blood meals taken from  hours, June 18 and 19.  In c o n t r a s t , two  simu-  (Cnephia minus) i n g e s t e d 48 and 50 m i c r o f i l a r i a e  t h i s b i r d on June 19, at 0430 hours.  this  from  In the evening of  June 19 the c a p t i v e grouse were p l a c e d o u t s i d e the r e s e a r c h cabin i n a shelter thenceforth.  where they experienced normal  On June 22, one C. minus. recovered at 0600  hours contained 6 m i c r o f i l a r i a e ,  and on June 25 two C. minus  and one S. aureum had i n g e s t e d 14, respectively, It  day-length  from t h i s grouse  16, and 28  microfilariae  at the evening  exposure.  i s suggested that the a r t i f i c i a l l y  Increased  day-length experienced by the c a p t i v e grouse #24 June s h i f t e d or a l t e r e d filarial normal  i n early  the p e r i o d of g r e a t e s t d a i l y micro-  abundance, and that when the grouse was  returned t o .  day-length the m i c r o f i l a r i a e resumed a temporal  d i s t r i b u t i o n s i m i l a r to that found during b i - h o u r l y in  testing  March. Most of the s p e c i e s of m i c r o f i l a r i a e  periodic  that show a  behavior reach t h e i r maximum numbers i n the p e r i -  pheral c i r c u l a t i o n nocturnally.  From b i r d s ,  these  C h a n d l e r e l l a c o l u m b i q a l l i n a e (Augustine, 1937)  in  include  131  ground-doves  (7), C h a n d l e r e l l a  bronzed g r a c k l e s 1929)  (43),  (?) Aproctoides  (Linstow, 1904) lissum  1964), E u f i l a r i a  Spiendidofilaria Hibler, (15).  1964,  caperata  (33)  and  Anderson (3)  filariae  lonoicaudata  (Chandler,  Hibler,  H i b l e r , 1964,  S.  1964,  picacardina  u n i d e n t i f i e d m i c r o f i l a r i a e i n crows  found, on the  other  hand, that the  of S p i e n d i d o f i l a r i a f a l l i s e n s i s  attained greatest  (Anderson,  (15), Robinson  able to completely and p e r i o d i c i t y of the  (48), and  r a p i d l y reverse  rupted  1). (42) were  (within two  days) the  avian m i c r o f i l a r i a e they  studied. exposed to con-  l i g h t , normal p e r i o d i c i t y of C. q u i s c a l i was and  the  conditions  than even the maxima recorded  of l i g h t  and  darkness.  to and  from the  small v e s s e l s  of the  Thurston  have demonstrated that t h e i r i n c r e a s e  lungs i s of the Hawking  (29)  lungs ( p a r t i c u l a r l y since Hawking  same magnitude as  regarded the migratory behavior of  conditions  i n the  and  i n the  t h e i r decrease p e r i p h e r a l l y ) .  a r i a e as a "compromise" between the favorable  periodic  i n numbers of . c i r c u l a t i n g m i c r o f i l a r i a e are  brought about by t h e i r r e g u l a r migrations  (30)  blood  under normal  It i s a g e n e r a l l y accepted view that the fluctuations  dis-  number of m i c r o f i l a r i a e i n p e r i p h e r a l  much g r e a t e r  and  Odetoyinbo  Odetoyinbo noted, as w e l l , that i n g r a c k l e s tinuous  micro-  1954)  abundance p e r i p h e r a l l y between 1200  hours i n the domestic duck ( t e s t e d February  Boughton e_t al  was  in  i n f r a n c o l i n s (28), S k r i a b i n o c t a s t r i a t o s p i c u l a  (Hibler,  1600  quiscali  microfil-  (presumably) more  lungs and  the  less  favorable  132 c o n d i t i o n s i n the p e r i p h e r y of the host's body, where, however, they  are r e a d i l y a c c e s s i b l e to t h e i r v e c t o r s .  that each m i c r o f i l a r i a lungs  has  He  postulated  an innate tendency to spend h a l f  its  time i n the  and h a l f i n the general  and  that t h i s innate rhythm i s modified  and  circulation,  synchronized  by  the 24-hour rhythm of the host's p h y s i o l o g y .  (This theory  fails  species such  to e x p l a i n the behavior  of non-periodic  as the u b i q u i t o u s S e t a r i a c e r v i Experimental  (Rudolphi,  a d m i n i s t r a t i o n of various s t i m u l i ,  exercise, anaesthesia,  and  for instance,  oxygen, anoxia, has produced  sometimes opposite responses by d i f f e r e n t filariae,  1819), however.)  i t likely  different,  s p e c i e s of  micro-  f o l l o w s that n a t u r a l - s t i m u l i evoke  equally diverse reactions. Most of the s t u d i e s on m i c r o f i l a r i a l have d e a l t with s p e c i e s i n f e c t i n g man, animals  maintained  one  a r t i f i c i a l illumination. has  considered  host.  as a r e s u l t  Perhaps f o r t h i s reason  no  (or micro-  on the c i r c a d i a n rhythm of the  l a t i t u d e of Vancouver the d a y l i g h t p e r i o d  i n c r e a s e s from 8 1/4 on June 21.  day-length,  might a l t e r p e r i o d i c i t y of  through i t s e f f e c t  At the  constant  that the normal d a i l y increment  decrement) i n day-length filariae  domestic mammals or  i n c a p t i v i t y - hosts which are f r e q u e n t l y  i n f l u e n c e d by a r e l a t i v e l y of  periodicity  hours on December 21 to 16 1/4  In view of the f i n d i n g s of Odetoyinbo  o t h e r s , the ease of r e v e r s i b i l i t y i n t e r r u p t i o n by constant  hours and  of p e r i o d i c i t y , i t s  illumination,  and the  concomitant  i n c r e a s e i n numbers of p e r i p h e r a l m i c r o f i l a r i a e ,  i s i t not  p o s s i b l e that m i c r o f i l a r i a l rhythm changes i n r e l a t i o n  133 to the host'a response Anderson had  i n June r a t h e r than February  he might have  between peak p e r i p h e r a l abundance  time of i t s s i m u l i i d Although  1961  1960,  and  vectors.)  the blood of grouse  m i c r o f i l a r a e m i a s of both Sk. i n October  (Perhaps i f fallisensis  c l o s e r synchrony  biting  daylight?  assessed the p e r i o d i c i t y of S.  microfilariae found  to extended  #25  flexivaginalis  showed moderate and Mf.  sp. B  smears made at 2-hour i n t e r v a l s i n March  contained so few of e i t h e r s p e c i e s that no c o n c l u s i o n  could be drawn r e g a r d i n g p e r i o d i c i t y . being t e s t e d t h i s grouse autopsy,  i t was  found  Three months a f t e r  d i e d of a s p e r g i l l o s i s .  that both  n e c r o t i c , probably the decrease  Since, at  lungs were e x t e n s i v e l y i n m i c r o f i l a r a e m i a s was  due  to the e f f e c t s of t h i s d i s e a s e . E.  Summary and Conclusions R e l a t i v e to Transmission The  o b s e r v a t i o n s presented i n t h i s s e c t i o n  suggest  that: 1) grouse  Transmission of Sk.  flexivaginalis  and Mf.  sp. B to  c h i c k s on Vancouver I s l a n d occurs a f t e r most c h i c k s  are at l e a s t  s e v e r a l weeks o l d ; and that t r a n s m i s s i o n of  S. p e c t o r a l i s to c h i c k s i n the i n t e r i o r of the province occurs a f t e r most of them are more than one month of 2)  Absence of Mf.  age.  p e c t o r a l i s i n f e c t i o n s from the hens  does not preclude t r a n s m i s s i o n of t h i s worm to her c h i c k s . 3)  U n l i k e i n f e c t i o n s with Sk.  flexivaginalis  sp. B, i n f e c t i o n s with S. p e c t o r a l i s c h i c k s than i n adult grouse, short-lived  (perhaps  due  and  Mf.  are more p r e v a l e n t i n  probably because the worms are  to immunity c o n f e r r e d by the  134 i n t e n s e response to t h e i r 4)  Unless  presence).  the v e c t o r s are present  i n f e c t i o n s i n c h i c k s probably f o r t r a n s m i s s i o n to other  during  c o n t r i b u t e very  grouse.  September, little  inoculum  135 III.  ARTIFICIAL TRANSMISSION  Frequent examination, of the p e r i p h e r a l blood grouse i n o c u l a t e d with supposedly i n f e c t i v e l a r v a e , and one  grouse i n t o which subadult  S. p e c t o r a l i s had  of a l l the  been t r a n s -  planted,  f a i l e d to demonstrate the  aemias.  When these b i r d s were s a c r i f i c e d , or died, between  3 weeks and  15 months a f t e r i n o c u l a t i o n , blood  harbored no m i c r o f i l a r i a e and t i s s u e was  out  At the  time most of t h i s experiment was  sp.  carried  aware that the m i c r o f i l a r i a e  (the source of inoculum) might be  than S. p e c t o r a l i s .  a c t u a l l y Mf.  but  not  lungs  the p e c t o r a l subcutaneous  of worms.  i n grouse #9  develop  from the  devoid  (1959) the w r i t e r was  other  anticipated m i c r o f i l a r -  of a  species  I f these m i c r o f i l a r i a e were  B, i t i s p o s s i b l e that some worms d i d  ( i n the unknown s i t e s of the  that m i c r o f i l a r a e m i a s  were very  adults of t h i s low  and  species)  very t r a n s i t o r y .  Anderson (3) succeeded i n producing i n f e c t i o n s with as few  as 68  l a r v a e ' o f .S. f a l l i s e n s i s  g e n e r a l l y used more larvae - up for  (?)Mf. sp.  B a greater  to 737  (although  per b i r d ) .  he  Perhaps  amount of inoculum i s r e q u i r e d  to produce patent i n f e c t i o n . The i n o c u l a t i o n had  flies  from which larvae were obtained  not been n a r c o t i z e d ,  and  that the process of i n j e c t i o n , which was was  i n j u r i o u s to the  larvae.  The  there was  c a r r i e d out  only other  occurs to the w r i t e r as a p o s s i b l e cause'of f a i l u r e was prevention  the  use  no  for evidence slowly,  f a c t o r that transmission  of grouse feed medicated at a l e v e l f o r  of b a c t e r i a l and  coccidial  disease.  136 IV. A,  DEVELOPMENT OF FILARIAL LARVAE  Fourth-Stage Larvae of S p i e n d i d o f i l a r i a p e c t o r a l i s n. sp. (Plate IV, f i g s . 4-7) A t o t a l of 21 f o u r t h - s t a g e larvae of S.  was  removed from under  D. o. p a l l i d u s  the p e c t o r a l s k i n of two  c h i c k s , J u l y 24, 1959.  pectoralis  sibling  Most of these  were about the same length - males, 0.97-1.2 mm;  larvae  females  1.0-1.2 mm,  and were probably a c q u i r e d at approximately the  same time.  A few, however, were somewhat l a r g e r or s m a l l e r ,  and l i k e l y r e p r e s e n t e d s l i g h t l y e a r l i e r or l a t e r  infections.  None of these l a r v a e evinced signs of an impending  moult  and  t h e r e f o r e i t i s probable that none had a t t a i n e d i t s maximum development tive  i n t h i s stage.  Since these worms, u n l i k e  infec-  larvae or f u l l y mature a d u l t s , had not reached a  comparatively s t a t i c  state r e l a t i v e  to growth,  measurements would give a m i s l e a d i n g p i c t u r e .  averaged Therefore,  the f i g u r e s presented i n Table XI are the'ranges recorded f o r each c h a r a c t e r and i n c l u d e a l l larvae The  cuticle  of' f o u r t h - s t a g e S. p e c t o r a l i s shows  delicate transverse s t r i a t i o n s . hypodermis  are w e l l developed.  ing c o n d i t i o n was chords comprise  found.  noted:  The  l a t e r a l chords of the  In young larvae the f o l l o w -  a n t e r i o r to the nerve r i n g  the  a narrow, g r a n u l a r , s y n c y t i a l band with  l o n g i t u d i n a l rows of i r r e g u l a r l y P o s t e r i o r to the nerve r i n g  alternating'large  two  nuclei.  2 l o n g i t u d i n a l rows of c u b o i d a l  c e l l s with small n u c l e i form between the aforementioned band and the c u t i c l e .  These rows are almost' contiguous and are  separated from each other by a very narrow s y n c y t i a l  strip  137 Table XI,  Range of measurements of f o u r t h - s t a g e l a r v a e of S. p e c t o r a l i s n. sp. ( i n microns unless s t a t e d o t h e r w i s e ) . Males 10  Number of Specimens Length  Females 11  0.750-2.50  (mm)  Distance to Nerve Ring  0.920-2.22 69-100  71-117  Distance to Excr. Pore  -  102-161  L a t e r a l Width of Head  24-33  27-31  D o r s o v e n t r a l W. of Head  22-32  21-31  W. half-way to Nerve Ring  26-32  25-32  Maximum Width  37-62  32-47  Width at Anus  3.1-49  -  283-541*  137-200  To Vulva or T e s t i s  Anlage  Length of Oesophagus Min.  '  231-402  Width of Oesophagus  ;  241-391  2-4  2-3  Max. Width of Oesophagus  10-21  7-17  Length of T a i l  49-77  60-105  T h i s . i n d i v i d u a l probably a t y p i c a l . i n c l u d i n g the longest, 283-402 .  Range f o r a l l others  138 containing  n u c l e i intermediate  a l t e r s considerably larvae the  lateral  as the  in size.  This  configuration  l a r v a develops, and  i n the  older  chords appear to comprise a double row  of  f u s i f o r m c e l l s bordered on each side by a syncytium of extremely numerous, very  small n u c l e i .  The  dorsal  and  v e n t r a l chords each c o n s i s t of a p a i r of separated  rows of  c e l l s with large n u c l e i , which p r o j e c t i n t o the body c a v i t y . The  arrangement of c e p h a l i c p a p i l l a e i s s i m i l a r to that  the  a d u l t , with one  l a t e r a l surface the  large p a p i l l a p r o j e c t i n g from each  at a c o n s i d e r a b l e  o r a l opening, and  o r a l opening.  distance,  a small p a p i l l a  resembles that of the  a d u l t , with a very  a n t e r i o r p o r t i o n the w a l l s  which i s much broader, i s very and  multinucleate.  the  i n t e s t i n a l epithelium end  At the  of the  The  l a r v a , bear  somewhat  oesophageo-intestinal extends a short  9  and  made up  The  the  narrow lumen,  and  epithelium  i n v e n t r a l view, about 18  The  p, broad,  bears a p a i r of t e r m i n o l a t e r a l f l e s h y protuberances those of the  intest-  rectum of both sexes i s short  of a small number of large c e l l s .  apex i s truncate  junction,  distance.forward  seems to c o n s i s t of a s i n g l e l a y e r of cuboidal inclusions.  vacuolate,  In most larvae  ine i s narrow, about 12 u. with a very  with g r a n u l a r  anucleate  a posterior portion  granular,  oesophagus.  the  oesophagus  narrow  of which, i n the  s t r i a t i o n s , and  from  on each side near the  o r a l opening than i n the adult c o n d i t i o n .  around the  about 14 | i ,  Amphids, however, are much c l o s e r to  numerous t r a n s v e r s e  of  a d u l t , each invaded by  a large  cell.  caudal and like  139 In observed, larged,  the arid  slightly the  the  male  cloacal  the  region  cloacal  protruding.  genital  larva,  tract  Even  extends  (which  i t  rectum.  In  anterior  the  larger  males  definitive of  these  pouches the in  structure  the  a  larger  a  the  near  of  to  the  vulva  vulvar extends by  an  a  mass  than  the  the  female  apparently,  rest,  large  larve,  the  body  the  The  a l l  the  t a i l  female  is  no  a  short  of  the  and  a  single  is  often  cuticle, the  the  anal  lips  posterior  larvae  developing  indication  of  by  a  digitiform  a  4  of  curved  distance  row  the  the  oesophageo-intestinal  Internally,  posterior  precursors  the  of  pore  body  enlarged,  to  and  cytoplasm.  the  abruptly  Within  visible  comprises  excretory  fourth-stage  there  adult.  a  is  any  ultimate  capped  dense  beyond  not  anterior  cuticle  and  is  cells  in  cells.  termination column  with  of  the  the  small  protrusion  Externally,  anterior  of  slight  In  of  reproductive  The  spicular  the  considerably  from  very  large  of  slightly.  lips  of  of  developed  suggestion  larger  junction.  pair  not  vague  region  is  had  the  formation.  larvae  proliferating  anus.  lumen  usually  the  protrude  of  The  examined  tract  is  of  cloacal  to  cuticle  portion  by  slightly)  of  a  are  oesophageo-intestinal  reproductive  of  en-  larva  layer  testular  indicated  a  not  posterior,  delicate  discernible.  In  the  number  is  much  the  spicules  outline  cell  smallest  was  noticeably  male  possess  small  is  the  indication  the  pore  the  spicular  rapidly  body  displaces  part  a l l  excretory  particularly  from  is  of  but  containing  pouches  in  sometimes  there  larvae,  of  lips,  junction  the  the  to  the  hyaline  backward 6  cells  "ovijectoral"  large core  bordered which  portion  are, of  the  140  vagina. lar  P o s t e r i o r to t h i s  column e x t e n d s  into  a short  or  as much as  in  a large  least  the  the  extends In  the  female,  little  cell  terminal  only  about  interior  of  separation lumen  of the cells,  most h i g h l y  of  of  and  haemocoele.  at  cuticle  the by  towards the  adjacent  rear  cells  of  the  appears  the  organ,  and  the  epithelium.  squamous  columns of  of  entire  squamous  the  terminating  epithelium  the  tract.  largest  to i n d i c a t e  formation.  of  Skriabinocta  flexivaqinalis  were f o u n d  Those  2 days  Shortly  later  still  i n the present  were u s u a l l y  a f t e r a t t a i n i n g the  migrate  t o the  in Culicoides  haemocoele  within  within  blood  alive  the  but  rather  of e n t e r i n g  the  haemocoele  the  t h o r a c i c m u s c l e s Where  develop-  commences. The  an  the  which  |i f e m a l e )  case  tract,  system,  parallel  cells 920  In t h e  p r e s u m a b l y were i n c a p a b l e  microfilariae ment  u. ( i n the  bifurcating  female) b e f o r e  i s covered  half-way the  75  cuboidal  U- l o n g .  2 cells  developed  ingestion.  meal more t h a n inactive,  20  massive m u l t i c e l l u -  caudad b e f o r e  mm  developed reproductive  Development  hours  as  2.2  about  Microfilariae 8  distance  \x ( i n the  exception  incipient  B.  520  ovoid  well  elongate In  a rather  2 p o s t e r i o r l y d i r e c t e d columns of  p a s s p o s t e r i o r l y f o r as  with  region  increase  length  first  external  i n width;  subsequently  t o 90  u or  l e s s and  particularly  near  the  excretory  cell  and  the  i n d i c a t i o n of the  increase  p o s t e r i o r end. rectal  cells  development  larvae  decrease  is in  f u r t h e r i n width, At  the  enlarge  same t i m e , and  become  the  141 h y a l i n e with large n u c l e i . reached c o n s i d e r a b l e  By the time- the sausage-stage i s  i n t e r n a l r e o r g a n i z a t i o n has occurred.  Advanced sausage-stage or f i r s t - s t a g e  larvae are about 160 p,  long with a maximum width of about 30 \x, and taper towards the a n t e r i o r end; the hypodermis i s w e l l and  a mass of p r o l i f e r a t i n g  cell  to the r e c t a l r e g i o n  tine; tail  a protruding  cells  slightly  developed  extending•from the e x c r e t o r y  i s probably the developing  intes-  c u t i c u l a r anal plug has formed and the  i s about 30 \x long, obtusely  c o n i c a l and lacks a c u t i -  c u l a r appendage. Larvae preparing not  to molt to the second stage were  observed nor were second-stage larvae found.  assumed that t h i s  stage i s of comparatively  Because of the d i f f i c u l t i e s larvae under f i e l d  conditons,  pected of. c o n t a i n i n g  It i s  short d u r a t i o n . .  involved  i n examing  many of the C u l i c o i d e s  third-stage  larvae were preserved i n  a l c o h o l f o r l a t e r d i s s e c t i o n i n the l a b o r a t o r y . the  third-stage  Unfortunately,  larvae removed from these f l i e s were u n s a t i s -  f a c t o r y f o r study; they f a i l e d and  sus-  to c l e a r w e l l i n g l y c e r i n e ,  because of t h e i r d e l i c a t e c u t i c l e  and now b r i t t l e  nature  they were u s u a l l y compressed and o f t e n broken on removal. The  f o l l o w i n g d e s c r i p t i o n of the ' t h i r d - s t a g e  i s based on 7 l i v i n g are i n microns. 50-69; d i s t a n c e  larvae  or h e a t - k i l l e d specimens; measurements  Length 448-513; d i s t a n c e to e x c r e t o r y  to nerve r i n g  pore 88-112; width of head 7-9;  maximum width 16-20; length of muscular oesophagus  76-(?)l37;  length of g l a n d u l a r  The head  oesophagus 54-81; t a i l  38-47.  bears 2 p a i r s of e q u a l l y w e l l developed p a p i l l a e and a p a i r  142 of s l i g h t l y p r o t r u d i n g head a s l i g h t  (?)amphids; immediately behind  c o n s t r i c t i o n occurs i n most specimens.  oesophagus of a l l specimens was tissue;  i t appeared that  followed  p a r t l y hidden by  p o s i t i o n to the  glandular  p o s t e r i o r r e g i o n was intestine. from the  The  oesophagus of the  difficult  s l i g h t l y elevated  a d u l t , but  anus.  abruptly  protuberances.  development of Sk.  flexivaqinalis  C u l i c o i d e s u n i c o l o r group) c l o s e l y p a r a l l e l s  Sk.  s t r i a t o s p i c u l a ( H i b l e r , 1964)  comparison of the 2 species  cent  are much longer not  be  At  longer.  The  glance,  flexivaqinalis  s t r i a t o s p i c u l a and  s u r p r i s i n g i f the t h i r d - s t a g e  i t would  larvae p a r t l y r e f l e c t e d  However, H i b l e r d i d not  B (see  been preserved.  s t a t e whether h i s  In t h i r d - s t a g e  larvae,  cent  longer  the  discrepancy  Mf.  specimens  than those g l y c e r i n e - c l e a r e d .  s i m i l a r shrinkage occurs i n S k r i abinocta  despite  larvae of  l a t e r ) the w r i t e r observed that l i v i n g  were about 20 per If  are  since he used "randomly s e l e c t e d specimens" probably h i s  m a t e r i a l had sp.  a  flexivaqinalis  measurements were from l i v i n g or g l y c e r i n e - c l e a r e d and  of  larvae of these  a d u l t s of Sk.  than those of Sk.  this difference.  first  of t h i r d - s t a g e  i n d i c a t e s that those of Sk.  almost 20 per  that  n. comb, i n C. haemato-  by H i b l e r (33). lengths  tail  terminates i n a  (in  potus as d e s c r i b e d  the  about 100 u,  Near i t s t i p the  c o n i c a l apex which lacks t e r m i n a l The  was  to d i s t i n g u i s h from the  broad i n t e s t i n e tapers  Comments.  granular  portion' comparable i n  curves s l i g h t l y dorsad or l a t e r a d and broadly  The  the narrow muscular r e g i o n  by a broader more granular  the  between the  larvae,  adult lengths  then,  there i s  143 not  l i k e l y much d i f f e r e n c e i n the s i z e of "the t h i r d - s t a g e  larvae of these 2 s p e c i e s . It i s perhaps t y p i c a l of t h i s genus that the of the t h i r d - s t a g e c o n d i t i o n was  larvae i s bent d o r s a l l y near i t s t i p .  This  noted i n the second-stage larvae of Sk.  striatospicula  by H i b l e r , and i t i s probably r e l a t e d to the  o f t e n bulbous caudal apex of the adults of both C.  tail  Development  of M i c r o f i l a r i a  (Plate VI, f i g s . It was  species.  sp. B i n S i m u l i i d a e  2-5)  p r e v i o u s l y pointed  been found to separate M i c r o f i l a r i a  out that no means have sp. B from Mf. p e c t o r a l i s .  Three of the grouse used f o r exposure o r i g i n a t e d i n the Okanagan V a l l e y , a r e g i o n where .S. p e c t o r a l i s occurs, the r e s t were captured on Vancouver I s l a n d . supposed that  I t was  originally  a l l of these grouse that harbored s h a r p - t a i l e d  m i c r o f i l a r i a e were i n f e c t e d with S.. pectora l i s .  However,  subsequent autopsies ,of a large number of grouse from Vancouver  I s l a n d without d i s c o v e r y  convinced the w r i t e r that  the m i c r o f i l a r i a e i n Vancouver  I s l a n d grouse must represent All  of adults of S. p e c t o r a l i s  another species  (Mf. sp. B).  grouse used f o r exposure were c a r e f u l l y examined when  they died or were s a c r i f i c e d , but i n none were S. p e c t o r a l i s adults  found.  In an attempt to determine whether  the c a p t i v e  grouse from the i n t e r i o r of the province had been host to S. p e c t o r a l i s or to Mf. sp, B, the t h i r d - s t a g e  larvae Which  developed i n s i m u l i i d s that had engorged on these grduse were compared with those from s i m u l i i d s that had engorged  on  144 grouse harboring  Mf. sp. B from Vancouver  Twenty-nine  Island.  larvae from the heads of s i m u l i i d s  which had fed on the c a p t i v e  adult Dendragapus o> p a l l i d u s  (#9, #15, and #49) and had been maintained upwards of 14 days were c l e a r e d i n g l y c e r i n e and compared with 28 s i m i l a r l y t r e a t e d larvae which had developed from microfilariae nosus,  of "species  B" harbored by 3 adult D. o. f u l i g i -  No r e a d i l y d e f i n a b l e  d i f f e r e n c e s were observed  between larvae o r i g i n a t i n g i n d i f f e r e n t host i n d i v i d u a l s or from d i f f e r e n t regions however, that  of the province.'  the i n t e s t i n e of t h i r d - s t a g e  I t was noted larvae o r i g i n a -  t i n g with D. o. p a l l i d u s hen #9 was c o n s i s t e n t l y , much narrower than the i n t e s t i n e of larvae developing from the Vancouver  I s l a n d grouse. Presumably t h i s d i f f e r e n c e i n i n t e s t i n a l  diameter was a f i x a t i o n a r t e f a c t , f o r larvae which had o r i g i n a t e d i n any of the c a p t i v e grouse always possessed a narrow i n t e s t i n e when examined alcohol-preserved  alive,' and larvae from  s i m u l i i d s which had fed on #9 showed  broad i n t e s t i n e s . Very l i k e l y  a l l 3 D. o. p a l l i d u s used f o r  exposure harbored Mf. sp. B.  However, the t h i r d - s t a g e  larvae of Mf. sp. B are r a t h e r s i m i l a r to the fourth-stage larvae of S. p e c t o r a l i s (see l a t e r ) , s i m i l a r to the t h i r d - s t a g e and S. p l c a c a r d i n a  (33).  and are extremely  larvae of S. f a l l i s e n s i s (4) Therefore,  i t may be very d i f f i -  c u l t t o d i s t i n g u i s h between the larvae of S. p e c t o r a l i s  145 and Mf.  sp. B, and one or more of the 3 D. p.. p a l l i d u s may  harbored Mf.  p e c t o r a l i s or both s p e c i e s of m i c r o f i l a r i a e .  Because of t h i s p o s s i b i l i t y from these 3 grouse  larvae which developed i n f l i e s  are omitted from the f o l l o w i n g  E a r l y develooment. abdominal  haemocoele w i t h i n 2 hours  site.  In Simulium  description.  M i c r o f i l a r i a e were found i n the after ingestion.  the body c a v i t y they proceed w i t h i n about tive  have  aureum t h i s s i t e  Once i n  a day to the appeared  defini-  to be the  f a t bodies a s s o c i a t e d with the ovary, whereas i n Cnephia minus i t seemed to be the r e t i c u l a r matrix of the ovary (Developmental  s i t e s were d i f f i c u l t  to determine  itself.  precisely  and i t i s p o s s i b l e that the f a t bodies are u t i l i z e d  i n both  hosts) . After attaining  t h i s s i t e the m i c r o f i l a r i a e  undergo  a gradual t r a n s i t i o n to the sausage-stage.  This process  requires  a considerable  1 to 3 days and i n v o l v e s i n i t i a l l y  r e d u c t i o n i n l e n g t h and an i n c r e a s e i n s i z e of the e x c r e t o r y and 4 r e c t a l  cells.  minimum l e n g t h , about  When the larvae have reached  100 \x, they are broad c e n t r a l l y  tapered towards each end. narrow elongate c u t i c u l a r cuticle  The  "tail",  14-17  u. long, which i s the  of the p o s t e r i o r p o r t i o n of the m i c r o f i l a r i a . of the  and p o s t e r i o r r e g i o n s and a f u r t h e r enlargement excretory  anterior  of the  cell. As the l a r v a e develop through the  drical.  and  caudal .extremity bears a  Subsequent development i n v o l v e s a broadening  (i.e.,  their  sausage-stage  e a r l y f i r s t - s t a g e ) they q u i c k l y become almost  cylin-  An a n v i l - s h a p e d , c u t i c u l a r anal plug forms and  146 e x t e n s i v e c e l l u l a r p r o l i f e r a t i o n i n t e r n a l l y produces developed hypodermal  l a y e r , and a m u l t i c e l l u l a r  a well  intestine  and s h o r t , narrow oesophagus, both l i n e d with d e l i c a t e cuticle.  The c e p h a l i c apex bears 4 minute,  refractile  c u t i c u l a r plaques c l u s t e r e d about the o r a l opening. body c u t i c l e remains  The  extremely t h i n , and the c u t i c u l a r  caudal appendage p e r s i s t s f o r the d u r a t i o n of the f i r s t stage without n o t i c e a b l e d i m i n u t i o n but always  reflexed  s h a r p l y along the d o r s a l surface of the p o s t e r i o r end. The smallest  sausage-stage  larva  stage from m i c r o f i l a r i a developed f i r s t - s t a g e  ( e x c l u s i v e of the t r a n s i t i o n a l  to larva) measured 125. (i; the best  l a r v a - 214 | i .  noted are ( i n m i c r o n s ) :  Other  dimensions  maximum width 24-38; d i s t a n c e to  e x c r e t o r y pore 34-39; and d i s t a n c e from anus to end of body proper 27-29.  One moulting f i r s t - s t a g e  l a r v a of 154 u.  had an oesophagus 34 u- long. Second-stage  larvae.  The youngest  l a r v a recovered had the f o l l o w i n g dimensions length 228; width of head  second-stage ( i n microns):  14; width at e x c r e t o r y pore 23;  maximum width 23; width at anus 20; d i s t a n c e to nerve r i n g 28, to 23.  e x c r e t o r y pore 48; length of oesophagus 58; length of t a i l A narrow, d e l i c a t e stoma about 6 U- long i s present i n  most second-stage to  lack n u c l e i .  l a r v a e ; the oesophagus i s narrow.and seems As the second stage progresses the oesophagus  elongates and i n some i n d i v i d u a l s vacuoles resembling those seen i n the oesophagus of adult S. p e c t o r a l i s  form i n i t s  p o s t e r i o r p o r t i o n ; the i n t e s t i n e becomes i n c r e a s i n g l y g r a n u l a r in  i t s a n t e r i o r r e g i o n and over most of i t s length r e d u c t i o n  147 in and  diameter occurs.  A small anal plug p e r s i s t s i n t h i s  the rectum transforms from a c l u s t e r of c e l l s i n t o a  r a t h e r t h i c k - w a l l e d s a c - l i k e s t r u c t u r e i n which the develops as a large vacuole.  The  caudal  c u t i c u l a r appendage which i s soon l o s t . i n c r e a s e s i n length the e x c r e t o r y r i n g becomes more prominent and p a i r of t e r m i n a l  caudal  As the  distance  the  next moult.  to nerve r i n g 44;  apical cuticle  i n d i c a t i v e of the  The  dimensions  as 312  Third-stage  larvae.  u., but  Table  morphometric d e s c r i p t i o n of the living  28.  pore  68;  A loosening  impending moult  t h e i r c u t i c l e when they were about 360  had  n o t i c e d when  ( i n microns):  to e x c r e t o r y  length of t a i l  observed i n larvae as short  based on 43  A  width of head 14; maximum width 3.4; width at  length of oesophagus 66; the  nerve  the c u t i c l e t h i c k e n s .  of a l a t e second-stage l a r v a were as f o l l o w s  anus 27;  larva  u. long; these subsequently enlarge •  the b r i e f p e r i o d before  length 344;  flap-like  a t r o p h i e s , the  protuberances were f i r s t  l a r v a e were about 250  during  cell  lumen  apex of the newly  emerged second-stage l a r v a bears a short, narrow,  the  stage  of  was  the m a j o r i t y  shed  u, l o n g . XII presents  a  t h i r d - s t a g e l a r v a of Mf.  sp. B  larvae from the heads of s i m u l i i d s which  fed on 5 c a p t i v e blue  grouse from Vancouver I s l a n d ,  and  28 g l y c e r i n e - c l e a r e d larvae from the heads of s i m u l i i d s which had  fed on 3 of the The  same grouse.  d i f f e r e n c e i n length between l i v i n g  glycerine-cleared  larvae r e p r e s e n t s  larvae c l e a r e d i n lactophenol averaging  409  u..  shrinkage.  showed l i t t l e  and  Fourteen  shrinkage,  148 Sixteen  l i v i n g t h i r d - s t a g e larvae from the abdomens  of s i m u l i i d s measured 370 The is  cuticle  smooth or, i n a few  transversely.  (345-406) | i .  of the t h i r d - s t a g e l a r v a of Mf. specimens, very d e l i c a t e l y  sp. B  striated  In en face view the stoma i s seen to be a  d o r s o v e n t r a l l y elongated  slit.  In d o r s o v e n t r a l view the  stoma appears to be bordered  l a t e r a l l y by s l i g h t  reminiscent  processes  of the dentiform  p a i r of what appear to be  prominences  of the A c u a r i i d a e .  large p a p i l l a e p r o j e c t , one  on each  s i d e , from the  l a t e r a l s u r f a c e s of the head.  view i t can be  seen that no i n t e r r u p t i o n s of the d o r s a l  v e n t r a l margins of the head occur is  "bullet-like".  visible  and  In  lateral  A short v e s t i b u l e , 1.5  only i n d o r s o v e n t r a l view and not  to 3 |i long, i n a l l specimens, In some  specimens the p o s t e r i o r part of the oesophagus i s up as broad, and  and  i t s form i n t h i s view  leads i n t o the narrow oesophagus, about 3 u- wide.  twice  A  appears more g r a n u l a r .  i n t e s t i n a l j u n c t i o n i s poorly demarcated.  The The  to  oesophageointestine is  very g r a n u l a r and moderately narrow, 6 to 9 (i, i n l i v i n g larvae. and  In most preserved  apparently  larvae the g r a n u l a r t i s s u e obscures  surrounds the i n t e s t i n e which seems to occupy  almost the e n t i r e diameter of the pseudocoele. ring  i s sometimes d e l i c a t e and  specimens c o n s i d e r a b l e  i n d i s t i n c t , but  cellular proliferation  o c c u r r i n g i n t h i s r e g i o n with This developing  connections  The  i n many seems to be  to the body w a l l  and  head.  the  a n t e r i o r part of the oesophagus g i v i n g i t an  of g r e a t e r breadth  nerve  t i s s u e o f t e n surrounds and  than i s a c t u a l l y the case.  obscures  appearance  The  excretory  149 Table X I I .  Measurements of t h i r d - s t a g e larvae of M i c r o f i l a r i a sp. B ( i n microns; mean followed by range i n parentheses).  Livina  To Nerve Ring To Excr.  -X-  Pore  Preserved l a r v a e , alvcerine-cleared  43  Number of Larvae Length  larvae  28  421  (308-478)  350  60  (53-69)'  46  (38-51)  78  (60-90)  77  (67-90)  ' 7  (6-9)  Lat. W. of Head ''  (295-430)  Max. W. of Body.  17  (16-22)  15  (12-18)  Width at Anus  14  (12-15)  12  (10-14)  8  (6-9)  W. of Caudal  Apex  L. of Oesophagus  80  (55-112)  89  (75-131)  Length of T a i l  29  (23-39)  26  (23-30)  52  (37-80)  69  (63-79)  Length of Rectum Gen. Primdrdium to Caudal Apex  83  (64-88); 110,142  ;  *Not seen i n a l l l a r v a e . 6 larvae and  only  4 larvae only. In 4 other larvae d e r i v e d from grouse #9 15, t h i s d i s t a n c e was 72, 74, 103 and 128 | i .  150 pore i s u s u a l l y v i s i b l e  at about the  i n t e s t i n a l j u n c t i o n , but the the  excretory  excretory  extends from the midventral The  pore a block l i n e to the  about 12 c e l l s ,  i n t e s t i n e , e i t h e r very  s l i g h t l y more a n t e r i o r to i t . region but at  cell  and  oesophageo-  the o u t l i n e of  Immediately  of about 12  i s short,  i n others  »  cells  a n t e r i o r part of  g e n i t a l primordium,.often d i f f i c u l t  d i s c e r n , comprising side of the  excretory  v e s i c l e are vaguely expressed.  p o s t e r i o r to the  intestine.  l e v e l of the  the  to  i s s i t u a t e d on the  ventral  near the r e c t a l r e g i o n  In some specimens the  i t i s elongated and  very  or  rectal narrow,  i n a l l i t expands to form a l a r g e , t h i c k - w a l l e d v e s i c l e the  apex.  anus, then narrowed, but The  caudal  tapered  t i p i s truncate  a p a i r of moderately large  little  towards the  i n v e n t r a l view and  bears  (up to 3 u. long) p a p i l l i f o r m  protuberances t e r m i n o l a t e r a l l y . D.  Influence  of Temperature on Development  Microfilaria the  larvae of Mf.  sp.  sp.  B.  The  B i n 2 periods  r a t e s of development of during  i n f e c t e d black  f l i e s were maintained at a mean temperature of  58  A f t e r 7 days only  f l i e s ; most of these larvae i n the  11 days a few  early third-stage  abdomen of a f l y ; and C. minus contained  abdomen.  days the  11 t h i r d - s t a g e  In c o n t r a s t ,  June  4,  larvae were found  sausage-stage; a f t e r  larvae were present  a f t e r 14.5  moulting second-stage l a r v a and i n the  first-stage  and  and  were compared.  i n the  20  spring  summer of 1961  (52-68-)°F.  Between May  the  only  i n the  remaining  larvae i n the head, and  2 moulting f i r s t - s t a g e  1  larvae  f l i e s maintained at a mean  151 temperature  of 74  (65-95)°F  between August 7 and August  y i e l d e d second-stage larvae a f t e r only 3.5 third-stage  flies  days and had  larvae i n t h e i r heads a f t e r 6.5 During the l a t t e r part of May,  days.  1961,  the few black  captured from i n f e c t e d grouse were maintained out-of-  doors as temperatures ranging between 52 and 68°F. mentioned  above,  normally, though  larvae of M i c r o f i l a r i a  As  sp. B developed  slowly, at these temperatures.  In June, to  e l i m i n a t e the adverse e f f e c t s of a n t - p r e d a t i o n and fly  15,  rainfall,  cages were moved onto the concrete f l o o r of the r e s e a r c h  cabin where the temperature e i t h e r side of i t s mean. weekly  f l u c t u a t e d only 4° at most, on  From e a r l y June to J u l y 7 the mean  temperature i n c r e a s e d g r a d u a l l y from 58 to 62°F.  t h i s p e r i o d no larvae of Mf.  sp. B developed beyond  During  the  sausage-stage i n the 75 black f l i e s  examined, and i n s e v e r a l  cases the l a r v a e were decomposing.  In a few f l i e s , captured  j u s t before the cages were brought i n d o o r s , and maintained f o r 2-3 weeks at the approximately constant temperature, most second- and t h i r d - s t a g e abdomen were dead.  larvae present i n the thorax' or  From J u l y 7 on, the cages were placed  outside the cabin f o r 3 to 6 hours each day where they were s u b j e c t to temperatures up to 95°F.  On J u l y 13, one C. minus  from J u l y 4 contained a c t i v e ,  l a t e second-stage  (temperature range J u l y 7-13,  57 to 94°F).  These  o b s e r v a t i o n s might  larvae  suggest that at moderately  low temperatures c o n s i d e r a b l e thermal f l u c t u a t i o n i s necessary for  s u s t a i n e d development,  (58°F)  l a r v a l development  since at the same mean temperature was  normal  i n those f l i e s  exposed to  152 a 16° range but was a r r e s t e d i n those c o n t i n u o u s l y exposed to a range of 8° or l e s s .  However, thermal constancy, per se.  e v i d e n t l y does not r e s t r i c t  filarial  development,  f o r Schacher  (49) found that Bruoia pahanoi developed normally at a constant 80°F, of  and H i b l e r  3 avian f i l a r i o i d s  (33) d e s c r i b e d normal  development  i n C u l i c o i d e s incubated at 85°F.  Furthermore, Hu (34) r e p o r t e d that the larvae of Wuchereria b a n c r o f t i r e q u i r e d an e x c e p t i o n a l l y long time - 33 days or more - but d i d a t t a i n t h i r d - s t a g e i n mosquitoes maintained at an almost constant 54.4°F at  (53.8-56.3);  a wider range of lower temperatures  either failed first  but i n mosquitoes (30-43°F)  kept  development  to occur or the larvae died a f t e r r e a c h i n g the  stage. If  the response to temperature  i s similar i n  Mf. sp. B and W. b a n c r o f t i then Mf. sp. B f a i l e d to develop s a t i s f a c t o r i l y under prolonged exposure to a narrow range of low temperatures because  these temperatures probably d i d not  a t t a i n the c r i t i c a l minimum l e v e l e a r l y enough. filaria  sp. B t h i s  l e v e l i s somewhat h i g h e r than f o r  W. b a n c r o f t i . and seems to be i n the v i c i n i t y S k r i abinocta f l e x i v a q i n a l i s . to  67 (54-83)°F  of t h i s s p e c i e s . only f i r s t - s t a g e  C u l i c o i d e s which had engorged  of 62°F.  I t was not p o s s i b l e  study i n d e t a i l the e f f e c t s of temperature  development  For M i c r o -  on the l a r v a l  However, i t was noted that at larvae were present i n 10 days e a r l i e r .  Third-stage  larvae were not found i n the heads of C u l i c o i d e s maintained at  68 (58-83)°F  before 17 days, but were present as e a r l y as  13 days when the temperature was 78 (50-97) F„ E.  Comments.on L a r v a l Development 1. General Among the species of Onchocercidae f o r which i t has  been s t u d i e d l a r v a l development f o l l o w s an almost p a t t e r n i n the i n v e r t e b r a t e host.  identical  The m i c r o f i l a r i a e penetrate  the w a l l s of the stomach or mid-gut, enter the abdominal haemocoele and invade an organ or t i s s u e which supports f u r t h e r development.  S e v e r a l developmental s i t e s have been  such as the abdominal haemocoele, the t h o r a c i c  reported,  skeletel  muscles, the abdominal, t h o r a c i c or c e p h a l i c f a t bodies, and the Malpighian restrictedly they reach exhibit  t u b u l e s ; the m i c r o f i l a r i a e of a species are  specific  as to the s i t e which they occupy.  their particular site  Once  t h e ' m i c r o f i l a r i a e begin t o  a decrease i n length and an i n c r e a s e i n width and c e l l  s i z e ; t h i s i s followed by a r e o r g a n i z a t i o n of i n t e r n a l s t r u c ture and i n c r e a s e d m i t o t i c a c t i v i t y  i n the s h o r t , s u b c y l i n -x-  drical  first-stage  l a r v a known as the sausage-stage.  Subsequent e l o n g a t i o n first  and c e l l u l a r p r o l i f e r a t i o n precedes the  moult, i n which the e x t e r n a l remnants of the micro-  filaria,  i t s c e p h a l i c and caudal  second-stage larvae i n c r e a s e s t r u c t u r a l e l a b o r a t i o n before  a p i c e s , disappear.  considerably  The  i n length and  the f i n a l moult i n the  This form i s apparently absent from the development of Dipetalonema v i t e (16), Dipetalonema mansonbahri (38) and Conispiculum g u i n d i e n s i s (45).  154 intermediate occurred and  host.  Very s h o r t l y a f t e r t h i s second moult  has  the t h i r d - s t a g e larvae leave t h e i r developmental  q u i c k l y migrate i n t o the head of the v e c t o r .  accumulate i n and v e r t e b r a t e host The  Here  around the mouthparts whence they  as the v e c t o r  site  they  enter  the  feeds.  r a t e of development i n the intermediate  host  appears to be g e n e t i c a l l y c o n t r o l l e d ; however i t i s i n f l u e n c e d somewhat by the degree of s u i t a b i l i t y major extent  by  The  of the host,  and to a  temperature. present  study demonstrates that the  larval  development of S k r j a b i n o c t a f l e x i v a g i n a l i s i n C u l i c o i d e s Microfilaria cercid  and  sp. B i n s i m u l i i d s conforms,to the b a s i c oncho-  pattern.  2. Development as an i n d i c a t o r of systematic p o s i t i o n . The  larvae of 4 other species of avian  have been f o l l o w e d process  i n Sk.  through t h e i r development.  f l e x i v a g i n a l i s and Mf.  onchocercids  Basically,  sp. B i s i d e n t i c a l  that d e s c r i b e d by Anderson (4) f o r S p l e n d i d o f i l a r i a s e n s i s i n s i m u l i i d s and by H i b l e r (33)  i n Culicoides»  to  falli-  for Skriabinocta  s t r i a t o s p i c u l a . S p l e n d i d o f i l a r i a p i c a c a r d i n a . and lonqicaudata  this  Eufilaria  There are, however, some f e a t u r e s  of morphology and developmental r a t e which vary w i t h i n t h i s group.  I t i s worth c o n s i d e r i n g whether these  features  r e l a t e d to p a r t i c u l a r adult and/or m i c r o f i l a r i a 1 and  i f so,.whether they  characters  can be used t o ' p r e d i c t the  p o s i t i o n of the unknown a d u l t s of M i c r o f i l a r i a  are  sp.  systematic B.  155  The  number of species  to permit extensive  s t u d i e d i s not  g e n e r a l i z a t i o n s to be drawn;  the data i n Table XIII do suggest the 1) The to n e i t h e r the the  discuss for  length of the  length  adult worms. the  of other  of mammals and  (It would be  species  of C a r d i o f i l a r i a  extent  by the  The  tall  attenuation  The  of the and  the  are g e n e r a l l y much  so i t may  be that extreme  length of the  third-stage  which have  third-stage  subterminal  first-stage  larval t a i l .  i s s l i g h t , however, and  lbnqicaudata  presence and  p o s i t i o n of the  while  adult  cloaca,  s i z e of a c u t i c u l a r appendage larva r e f l e c t  This i s to be  the degree of tail.  The  independent  expected since  s t r u c t u r e i s the remnant of the m i c r o f i l a r i a l which v a r i e s c o n s i d e r a b l y  The  i s the' s h o r t e s t i t s length  s t r u c t u r e seems to be q u i t e  affinities.  very  i s f o r e c a s t to some  acuteness of the m i c r o f i l a r i a l  occurrence of t h i s systematic  perhaps be noted that  and  are p a r a s i t e s of b i r d s ) .  length of the  s c a r c e l y p r e d i c t s the  on the  seem to support t h i s  length of adult t a i l  of l a r v a l E.  3)  the data a v a i l a b l e  (Pseudaproctella).  v a r i a t i o n among a l l 6 species the t a i l  of  of i n t e r e s t to study the development of  long m i c r o f i l a r i a e and 2)  length  t h e i r adults and m i c r o f i l a r i a e  d i f f e r e n c e s are r e f l e c t e d i n the larva.  larva i s related  vertebrates  I t should  as w e l l , and  inferences:  nor the  larvae of non-avian f i l a r i o i d s  o f t e n longer  nevertheless  the w r i t e r ' s i n t e n t i o n to  reptiles  longer than those of b i r d s and are  third-stage  While i t i s not  succeeding i n f e r e n c e s . third-stage  following  of the m i c r o f i l a r i a  filarioids  filariae  large enough  caudal  between c l o s e l y r e l a t e d  of  this extremity  species.  Table X I I I .  Some developmental and d e f i n i t i v e f e a t u r e s of 6 avian onchocercids.*  flex.  s t r i a t.  fallis.  B.  picacard.  lonqic.  9  15 76 36 110 mod.  8 v.short 16 60 pres.  Adults L. male (mm) L. male t a i l {\x) L. female (mm) L. female t a i l (mm) Terminal protub.  13 89 30 240 abs.  5 61 14 68 abs.  12 65 32 68 pres.  170 )lunt  154 blunt  105 bl.-pt.  abs. 384** .42 abs.  abs. 397 30 abs.  short 432 -30 large  long 350 26 large  13-78° 17-68°  11-85° 21-67°  7-62° 10-58°  6.5-74° 11-58°  ? ?  ? ?  Microfilariae Length - (ji) Caudal apex  ;  215 sharp-pt.  127 bl.-pt.  98 sharp-pt  Larvae .. l s t - s t . caud. append.L. 3 r d - s t . -larva ( u . ) L. 3 r d - s t . t a i l Terminal protub. Time to reach head (days at temp, i n F )  abs. 505 38 mod.  long 537 1.9 pres.  8-85° 12-67°  7-85° 10-67°  * These data are from the f o l l o w i n g sources: S k r j a b i n o c t a f l e x i v a g i n a l i s and Mf. sp. B - t h i s study; S p l e n d i d o f i l a r i a f a l l i s e n s i s - Anderson (3, 4 ) ; .S. p i c a c a r d i n a . S k r i a b i n o c t a s t r i a t o s p i c u l a . and E u f i l a r i a longicaudata H i b l e r (33). —  ** Length of unpreserved larvae reduced by 20% to correspond glycerine-clearing i n the other s p e c i e s .  to the e f f e c t of  157 (All  known s p e c i e s  microfilariae; first-stage  of S k r i a b i n o c t a produce b l u n t - t a i l e d  i t i s probably a reasonable p r e d i c t i o n that the  larvae of a l l these species 4)  Terminal caudal  lack  t h i s appendage)=  protuberances i n the t h i r d - s t a g e  l a r v i pr§§ag§ t h o i r prgsence i n the a d u l t .  I t Is not c i g a r  whether the protuberances of l a r v a l and adult Eufilaria  lonoicaudata  are homologues nor whether they are  homologous t o those i n S p l e n d i d o f i l a r i a . r e l a t i o n s h i p obtains  only w i t h i n generic  converse r e l a t i o n s h i p i s l i k e l y  Possibly t h i s limits  (though the  to be more u n i v e r s a l l y true.)  5) C l o s e l y r e l a t e d s p e c i e s developmental times.  (male)  are s i m i l a r i n t h e i r  The converse i s probably not t r u e , f o r  many f i l a r i o i d s  i n d i f f e r e n t genera develop at approximately  the  P r e c i s e comparative s t u d i e s on the e f f e c t s of  same r a t e .  different  constant  temperatures on r a t e of development of  organ systems and on d u r a t i o n  of each l a r v a l stage are long i  overdue. materially  Conceivably,  such s t u d i e s could  contribute  to a b e t t e r understanding of f i l a r i a l  interrela-  tionships. 6) On the b a s i s of some of the foregoing the w r i t e r suggests that the adults of M i c r o f i l a r i a probably belong to a species  designated  sp. B  of S p l e n d i d o f i l a r i a with a  moderately short t a i l which bears a p a i r of t e r m i n a l protuberances.  inferences  caudal  I t i s worth r e c a l l i n g that a s i n g l e male  as S p l e n d i d o f i l a r i a sp.- A was taken from a grouse  which harbored Mf. sp. B.  The c h a r a c t e r s  i n f e r r e d f o r the  unknown a d u l t s of Mf. sp. B c o i n c i d e w e l l with those of Splendidofilaria  sp. A as d e s c r i b e d  on pp. 27-28.  158 V. A.  ORNITHOPHILIC DIPTERANS  Haematophagous Dipterans C o l l e c t e d from the Study Areas The s p e c i e s l i s t e d  i n Appendix V I I I are those  captured i n both study areas from exposed grouse, from  man,  or from net-sweepings  section,  i n the v i c i n i t y of man.  In t h i s  only those d i p t e r a n s recovered from grouse w i l l be  considered.  I d e n t i f i c a t i o n s were made with the a i d of keys from v a r i o u s sources and i n f o r m a t i o n s u p p l i e d by d i p t e r o l o g i s t s .  Determina-  t i o n s f o r the w r i t e r ' s specimens of C u l i c o i d e s were confirmed and c o r r e c t e d by Mr. who  J . A. Downes, Canada Dept. of A g r i c u l t u r e ,  pointed out (1964, i n l i t t . )  that  "the whole genus  C u l i c o i d e s i s i n a very u n s a t i s f a c t o r y s t a t e , and  ... the  c o a s t a l B. C. forms i n p a r t i c u l a r have r a r e l y been  collected",  and that most of the s p e c i f i c names cover " . . . q u i t e a multitude of c l o s e l y r e l a t e d Undoubtedly Simuliidae,  forms..."  a s i m i l a r s i t u a t i o n e x i s t s i n the  f o r a small c o l l e c t i o n of black f l y l a r v a e , pupae,  and a d u l t s from the U n i v e r s i t y of B r i t i s h Columbia campus, which I  submitted i n 1961  to Dr. K. R o t h f e l s , U n i v e r s i t y of  Toronto, comprised 3 undescribed species and 2 new occurrence i n Canada.  records of  U n f o r t u n a t e l y , no r e p l i e s have been  r e c e i v e d i n answer to my r e q u e s t s f o r i d e n t i f i c a t i o n of a d u l t s , pupae, and larvae c o l l e c t e d i n the study areas and submitted to s e v e r a l a u t h o r i t i e s .  Consequently, some of the  determinations of the s i m u l i i d s recovered are t e n t a t i v e . s t a t u s of the abundant,  small dark Cnephia c o l l e c t e d  grouse i s open to q u e s t i o n ,  f o r there are i n d i c a t i o n s  2 very s i m i l a r forms are i n v o l v e d here.  Because  from that  of the  The  159 difficulty  i n d i s t i n g u i s h i n g between these forms, and because  no d i f f e r e n c e was noted i n t h e i r s u i t a b i l i t i e s  as intermediate  h o s t s , they are considered here as Cnephia minus (D. and S.), which they c l o s e l y  resemble.  The f o l l o w i n g d i p t e r a n s were c o l l e c t e d from grouse, g e n e r a l l y from those b i r d s exposed f o r the purpose of attracting  insects.  Culicidae Cu1iseta i n c i d e n s Mansonia  (Thorn.) - Nanaimo Lakes  perturbans (Walk.) - Nanaimo Lakes  Ceratopogonidae C u l i c o i d e s c r e p u s c u l a r i s M a l l , group - Trout Creek and Nanaimo Lakes Culicoides unicolor  (Coq.) group - Trout Creek and Nanaimo Lakes  C u l i c o i d e s sp.  - Trout Creek  C u l i c o i d e s sp. 1  - Nanaimo Lakes  Simuliidae Prosimulium dicentum D.  and S. - Trout Creek and Nanaimo Lakes Prosimulium p l e u r a l e M a l l . - Trout Creek and Nanaimo Lakes Simulium  (Eusimulium) aureum F r i e s - Trout Creek and Nanaimo Lakes  Simulium  (Eu.) ?osborni  (Stains and Knowlton)  - Trout Creek  Simul.ium (Simulium) canadense Hearle - Trout Creek S. (S,) o c c i d e n t a l e Towns. - Nanaimo Lakes S. (S,) venustum Say - Nanaimo Lakes Cnephia minus (D. and S.) - Trout Creek and Nanaimo Lakes  160 S,  (Eu.) sp.  - Trout Creek  S.  (Eu.) sp.  - Trout Creek  Hippoboscidae Ornithoica vicina Ornithomvia  frinoillina  Neolipootena B.  (Walk.) - Quinsam, Vancouver I s .  ferrisi  L a t r . - Quinsam, Vancouver I s .  (Beq.) - Quinsam  G r o u s e - B i t i n g I n s e c t s as Intermediate Hosts and V e c t o r s In  this  s e c t i o n the i n s e c t s c o l l e c t e d  on the grouse range w i l l  from grouse or  be considered from the standpoint of  t h e i r p o t e n t i a l as v e c t o r s of the f i l a r i a e  of grouse.  1. Mallophaga Biting  l i c e were present on s e v e r a l of the adult  blue grouse captured and maintained i n c a p t i v i t y . the  Because of  close p r o x i m i t y of the cages hand-reared grouse soon became  infested  as w e l l .  The adult grouse harbored M i c r o f i l a r i a  sp. B  and Mf. f l e x i v a q i n a l i s i n t h e i r blood, but no i n f e c t i o n s developed i n the young development  of " F i l a r i a  grouse. cvpseli"  reconditum (of dogs), i n b i t i n g Heterodoxus  Although Nelson (39) obtained (of s w i f t s ) , and Dipetalonema l i c e , Dennysus h i r u n d i s and  s p i n i q e r . r e s p e c t i v e l y , there i s no reason to  suspect that the f i l a r i o i d s  of grouse are t r a n s m i t t e d by  mallophagans. 2. Tabanidae While the r o l e of Chrysops s i l i c e a c e a mediate host f o r Loa loa of man Bennett  as an i n t e r -  i s w e l l known, and although  ( l l ) obtained a few specimens of Chrysops spp. from  raven and crow i n O n t a r i o , none of the s e v e r a l species of  161 tabanids observed i n the B r i t i s h Columbia  study areas was  n o t i c e a b l y a t t r a c t e d to grouse. 3. C u l i c i d a e Of the 2 species of mosquitoes exposed  a t t r a c t e d to grouse  near a small marsh at Nanaimo Lakes only  perturbans was captured engorged.  Mansonia  Mosquitoes of 4 other  s p e c i e s a t t r a c t e d t o man and placed with a grouse i n the collecting  cage o v e r n i g h t , f a i l e d  to feed on the b i r d .  S i x t e e n female M. perturbans that had taken blood meals c o n t a i n i n g Mf. sp. B from grouse #4, #24, (and #49) i n l a t e J u l y y i e l d e d only dead m i c r o f i l a r i a e when d i s s e c t e d 2 t o 18 days l a t e r .  Although mosquitoes  transmit many f i l a r i o i d s of  mammals and r e p t i l e s they are not w e l l equipped f o r b i t i n g birds.  Culex f a t i q a n s was i n c r i m i n a t e d as a v e c t o r f o r  (?)Aproctoides l i s s u m of an Indian f r a n c o l i n  (44); however  there i s no evidence to suggest that mosquitoes filarioids  of g a l l i n a c e o u s b i r d s i n B r i t i s h  transmit the  Columbia.  4. Hippoboscidae Bequaert  (14) r e p o r t e d Ornithomvia f r i n q i l l i n a  from  blue, r u f f e d , and F r a n k l i n ' s grouse i n the i n t e r i o r of B r i t i s h Columbia,  and O r n i t h o i c a v i c i n a . which he c o n s i d e r e d a c c i -  d e n t a l , from blue grouse on Vancouver found no louse f l i e s  Island.  Bennett (12)  on 54 r u f f e d grouse and 9 spruce grouse  i n Algonquin Park, O n t a r i o , but the w r i t e r and others have  Except f o r one which had a 1.9 mm t h i r d - s t a g e l a r v a very s i m i l a r t o that of S e t a r i a spp. i n i t s labium.  162 c o l l e c t e d 0. f r i n q i l l i n a Vancouver  o c c a s i o n a l l y from blue grouse on  I s l a n d , and l e s s f r e q u e n t l y 0. v i c i n a  Neolipoptena  ferrisi.  and  Since the n a t u r a l hosts of the  s p e c i e s are deer i t i s h i g h l y u n l i k e l y to' transmit of  grouse.  last  filarioids  A l l of s e v e r a l dozen male N. f e r r i s i allowed to  remain overnight with a hooded grouse i n the c o l l e c t i n g were r e c o v e r e d the next morning, Nelson  unfed, and many of them dead.  (40) showed that Hippobosca  Dipetalonema  lonqipennis  mediate hosts f o r the f i l a r i o i d s 0. f r i n q i l l i n a  and one  grouse h a r b o r i n g Mf.  seldom until  transmitted  d r a c u n c u l o i d e s of dogs, but f o r the f o l l o w i n g  reasons i t i s improbable that avian louse f l i e s  larvae,  cage  of grouse:  female 0. v i c i n a  are i n t e r -  1") one  female  from a y e a r l i n g blue  f l e x i v a g i n a l i s contained no developing  2) the more abundant found on grouse,  s p e c i e s , 0. f r i n g i l l i n a . i s  3) louse f l y p o p u l a t i o n s • a r e low  J u l y and August, and by t h i s time grouse c h i c k s are  no longer brooded by the hens (any p o s s i b l e interchange of f l i e s by contact of hosts has been reduced), and the cock birds  ( p o t e n t i a l source of inoculum) have begun to l e a v e the  summer range, and 0. f r i n g i l l i n a  4) Bennett- (12) has shown that  "...rarely  female  leave a good host u n l e s s they are  disturbed." 5. S i m u l i i d a e  as hosts f o r Mf.  sp. B  Simulium aureum and Cnephia minus were by f a r the most abundant of  the other 8 s p e c i e s was  viduals. are  s i m u l i i d s c o l l e c t e d from exposed  grouse.  Each  r e p r e s e n t e d by only 1 or 2 i n d i -  I f the numbers of f l i e s  a true i n d i c a t i o n of r e l a t i v e  captured from exposed  grouse  abundance, then among the  163 s i m u l i i d s , only S.  aureum and C. minus can be  p o t e n t i a l l y e f f e c t i v e v e c t o r s of f i l a r i o i d s It was captured  found that M i c r o f i l a r i a  on Vancouver I s l a n d (#4,  #8,  considered  of grouse.  sp. B from the  #24,  #25,  and #33)  loped to t h i r d - s t a g e larvae i n both Simulium aureum Cnephia minus. supports  The  grouse  a v a i l a b l e i n f o r m a t i o n , presented  deve-  and earlier,  a view that both groups of grouse harbored M i c r o f i l a r i a  sp. B but s i n c e t h i s  conjecture  cannot be v e r i f i e d ,  the  of d i s s e c t i n g f l i e s which b i t Okanagan grouse w i l l not  results be  considered. I n g e s t i o n of m i c r o f i l a r i a e .  Bennett  (13) , using  32 P  i n ducks and  a hen,  only the  l a r g e s t , S.  aureum.  In the present  found that of 6 o r n i t h o p h i l i c s i m u l i i d s  c r o x t o n i . took up more blood than study,  p r e c i s e volumetric  t i o n s could not be made; however, i t was blood meal of C. minus was aureum.  I t i s not  S.  determina-  observed that  the  u s u a l l y s m a l l e r than that of  S.  known whether t h i s i n d i c a t e s that S. aureum  u s u a l l y engorges to i t s maximum c a p a c i t y while  C. minus does  not, or that they both engorge f u l l y but C. minus i s p h y s i c a l l y incapable  of i n g e s t i n g as much blood.  s m a l l e r f l y the The  Since C. -minus i s a  l a t t e r i n t e r p r e t a t i o n may number of Mf.  engorged C. minus and minutes a f t e r these  be more a p p l i c a b l e .  sp. B i n the blood- meals of  9 engorged S. aureum was  f l i e s had  recorded  15 a  few  fed on s e v e r a l i n f e c t e d grouse.  Although l e v e l s of m i c r o f i l a r a e m i a s v a r i e d c o n s i d e r a b l y among these  grouse i t was  of about twice same b i r d .  found that S. aureum had  as many m i c r o f i l a r i a e  as had  i n g e s t e d an average C. minus from the  164 Suitability  f o r development.  The only i n d i v i d u a l s of  S, aureum and C. minus c o n s i d e r e d here were those t o which a l l the  following conditions applied:  Vancouver  I s l a n d blue grouse,  l ) obtained engorged  from  2) obtained from grouse with  s u f f i c i e n t l y high m i c r o f i l a r a e m i a s of Mf. sp. B that C. minus was known to i n g e s t  a minimum of 3 m i c r o f i l a r i a e ,  s p e c i e s of s i m u l i i d s c o l l e c t e d  3) both  from the same b i r d "at approxi-  mately the same time of day, and  4) d a i l y exposure to the  p r e v a i l i n g outdoor maximum temperatures, f o r at l e a s t 9 days a f t e r capture. D i s s e c t i o n of these f l i e s #33) y i e l d e d the f o l l o w i n g r e s u l t s :  (from grouse #4, #8, and l ) of 57 C. minus. 8  contained l a r v a e , with 7.3 (1-16) larvae per p o s i t i v e f l y (or 1.0 larvae per f l y ,  for a l l f l i e s ) ;  2) of 14 S. aureum. 11  contained l a r v a e , with 30 (3-170) larvae per p o s i t i v e f l y (or 24 larvae per f l y ,  for a l l f l i e s ) .  The number of f l i e s  from  each b i r d i s not large enough to permit very meaningful comp a r i s o n between C. minus and S. aureum f o r i n d i v i d u a l grouse. Howe,ver, i t i s worth mentioning that the 4 p o s i t i v e C.  of 11  minus from grouse #8 (with a high m i c r o f i l a r a e m i a )  con-  t a i n e d 3, 15, 15, and 16 l a r v a e , whereas the 2 S. aureum from t h i s b i r d contained 48 and 170 l a r v a e . Since S. aureum i n g e s t s about twice as many microfilariae  per blood meal as C. minus the above data suggest that  a much g r e a t e r p r o p o r t i o n of the i n g e s t e d m i c r o f i l a r i a e (Mf. sp.  B) develop s u c c e s s f u l l y i n S. aureum. Abundance and d i s t r i b u t i o n .  of  During the major  period  abundance s i m u l i i d s were approximately twice as numerous  165 at Trout Creek i n the i n t e r i o r oif the p r o v i n c e , 'as at Nanaimo Lakes on Vancouver  Island.  In the f i r s t  l o c a l i t y 52 a r b o r e a l  exposures, averaging 14 minutes each, y i e l d e d per exposure  10.6 s i m u l i i d s  i n 1959, while at Nanaimo Lakes 46 a r b o r e a l  exposures averaging 15 minutes, y i e l d e d 4.5 s i m u l i i d s per exposure i n 1962.  At Trout Creek S. aureum a n d C . minus were  c o l l e c t e d from grouse i n the r a t i o of 4:1, whereas at Nanaimo Lakes C. minus was approximately 5 times as abundant  as S.  aureum. 6. C u l i c o i d e s as hosts f o r S k r i a b i n o c t a I n g e s t i o n of m i c r o f i l a r i a e . coides c o l l e c t e d  flexivaginalis  In most of the C u l i -  from c a p t i v e grouse h a r b o r i n g Mf. f l e x i v a o i n a l i s  no d e v e l o p i n g l a r v a e were found..  Many C u l i c o i d e s examined  s h o r t l y a f t e r feeding on these b i r d s contained no m i c r o f i l a r i a e , but the m i c r o f i l a r a e m i a s of these b i r d s were low. bable that Mf. f l e x i v a o i n a l i s was not abundant  I t i s pro-  enough i n t h e i r  p e r i p h e r a l blood to be i n g e s t e d i n the small blood meal taken by C u l i c o i d e s .  Grouse #3 showed an extremely high monotypic  m i c r o f i l a r a e m i a of Sk. f l e x i v a g i n a l i s  (180 m i c r o f i l a r i a e  ina  large drop of p e r i p h e r a l blood as compared to 3-50 m i c r o f i l a r i a e of t h i s  species i n a s i m i l a r volume of blood from the other  grouse used).  F i f t e e n C u l i c o i d e s examined s h o r t l y a f t e r capture  from t h i s grouse contained 245 m i c r o f i l a r i a e , an average of 16.  Even though Sk. f l e x i v a o i n a l i s does not develop i n the  s i m u l i i d s the r e l a t i v e  s i z e of the blood meals of C u l i c o i d e s  and s i m u l i i d s i s pointed up by the d i s c o v e r y that 4 Cnephia minus c o l l e c t e d microfilariae,  at the same time from grouse #3, contained 262 an average of 65.  166 Suitability  f o r development.  were found i n a large number of C. C u l i c o i d e s sp, harboring  Mf.  1 which had  no developing a few  1 ingested  group, both i n the Island. host  of the  f o r Sk.  the  other  hand, development  l e a s t abundant form, C.  f l e x i v a q i n a l i s . the c r e p u s c u l a r i s and  and  on Vancouver intermediate  w r i t e r r e f r a i n s from conC.  sp.  1 are  i n most of the  unsuitable  grouse exposed  distribution.  B i t i n g midges were  much more abundant at Nanaimo Lakes than at Trout first  area  3.3  whereas at Trout sure i n 1959, the  unicolor  low. Abundance and  the  species  c r e p u s c u l a r i s group y i e l d e d  i n t e r i o r of the province  because the m i c r o f i l a r a e m i a s  from grouse  in this  While C. u n i c o l o r group i s a s u i t a b l e  c l u d i n g that C.  were  On  larvae  maintained i n c a p t i v i t y ,  larvae were found; C.  i n most f l i e s  engorged  few m i c r o f i l a r i a e and  sausage-stage l a r v a e .  occurred  third-stage  c r e p u s c u l a r i s group or  been captured  f l e x i v a q i n a l i s and  C u l i c o i d e s sp.  No  C u l i c o i d e s were c o l l e c t e d per  Creek 0.12  and  1.1  yellowish (C. sp.  species  l ) ; C.  as o f t e n , and numbers.  So  c r e p u s c u l a r i s group was C. u n i c o l o r group was  few  per  expo-  On Vancouver I s l a n d a small,  plain,  f o r which no name i s p r e s e n t l y a v a i l a b l e , encountered about h a l f  present  i n moderately  C u l i c o i d e s were c o l l e c t e d at Trout  that species  composition was  seemed to be  C.  group.  C u l i c o i d e s was  In  exposure,  C u l i c o i d e s were captured  per exposure i n 1960.  p r i n c i p a l grouse-biting  Creek.  Creek  not determined, however, some  c r e p u s c u l a r i s group and  others  C.  unicolor  low  167  7. Summary Ten s i m u l i i d s , 4 C u l i c o i d e s . 3 h i p p o b o s c i d s , c u l i c i d were found of these  are  most are not  a t t a c k i n g grouse i n the  study  and 1  areas.  incapable of supporting l a r v a l development  Some and  abundant enough as g r o u s e - b i t e r s • t o play a s i g n i -  f i c a n t r o l e i n the t r a n s m i s s i o n of f i l a r i o i d s  of t e t r a o n i d s .  Both C. minus and S. aureum are s a t i s f a c t o r y intermediate f o r Mf,  sp. B, and because of t h e i r abundance are probably  v e c t o r s of t h i s f i l a r i o i d .  and  suffi-  c i e n t l y abundant that i t should be considered a v e c t o r . cannot be concluded  the  C u l i c o i d e s u n i c o l o r group i s an  adequate i n t e r m e d i a t e host f o r .Sk, f l e x i v a q i n a l i s  it  hosts  Because  p o s i t i v e l y t h a t the other, more abundant  s p e c i e s of C u l i c o i d e s are u n s u i t a b l e as intermediate hosts f o r Sk.  f l e x i v a q i n a l i s and because s p e c i f i c determinations were  o f t e n u n c e r t a i n , the species of C u l i c o i d e s captured are pooled  from grouse  i n the subsequent p r e s e n t a t i o n s of behavior  of  vectors. C.  I n f l u e n c e of Exposure Duration on C o l l e c t i o n Other workers  minutes.  E a r l y i n the present  sures of t h i s  i n any  55)  expqsed b i r d s fox-. 20 to 30_  study i.t was  found  that expo-  length r e s u l t e d i n the l o s s of some' of the  engorged f l i e s . tance  (ll,  Efficiency  Since exposure techniques  are o f major impor-  study of the v e c t o r s of avian haematozoa  s e c t i o n i s devoted  to an assessment of the r e l a t i v e  of exposures of d i f f e r e n t d u r a t i o n .  this  efficiency  168 1. C o l l e c t i o n of Table XIV.  simuliids  C o l l e c t i o n of s i m u l i i d s from a r b o r e a l evening exposures of d i f f e r e n t d u r a t i o n at the same s i t e i n three y e a r s . *  (1)  (2)  121  1961  26  265  204  10  7.8  1.3  1962  18  260  165  15  9.2  1.7  1964**  18  540  174  30  9.7  3.1  (2)/(l)  (2)/(3)  (3)/(l)  Year  * Based on the 6 most productive #8 Nanaimo Lakes.  evenings during June at s i t e  S  ** Basic data are from Woo Explanation (1) (2) (3) (2) / ( l ) (3) / ( l ) (2)/(3)  (55).  of columns: Number of exposures. T o t a l exposure time ( i n minutes). T o t a l number of s i m u l i i d s captured. Average exposure d u r a t i o n ( i n minutes). Average number of s i m u l i i d s per exposure. T h e o r e t i c a l time r e q u i r e d to capture one s i m u l i i d ( i n minutes). For purposes of comparison of c o l l e c t i o n  it  i s assumed that the p o p u l a t i o n d e n s i t y of b i t i n g  approximately all  efficiencies flies  the same during the p e r i o d of peak abundance i n  3 years. S e v e r a l t e n t a t i v e c o n c l u s i o n s r e g a r d i n g the  ship between exposure d u r a t i o n and Table XIV.  captures  are suggested  relationby  l ) I n c r e a s i n g the length of exposure r e s u l t s i n  o b t a i n i n g more s i m u l i i d s i n each exposure.  Probably  there i s  a maximum exposure d u r a t i o n at which e q u i l i b r i u m between and  was  departing simuliids  would be taken  i s reached  i n approximately  and beyond which  constant  numbers.  arriving  simuliids This i s borne  169 out  by the i n d i c a t i o n that  2) E f f i c i e n c y decreases i n a p p r o x i -  mately d i r e c t r e l a t i o n to i n c r e a s e d exposure the  From  p o i n t of view of time expended i t i s obvious that 30-minute  exposures  are not s u f f i c i e n t l y more p r o d u c t i v e than 10- or  15-minute exposures for  duration.  flies  to j u s t i f y w a i t i n g 2 to 3 times as long  to b i t e the b i r d .  Apart from t e c h n i c a l c o n s i d e r a -  t i o n s , i t must be kept i n mind that the longer the  exposure,  the  i n t o the  g r e a t e r the danger  of r e l e a s i n g i n f e c t e d f l i e s  vicinity. Another element which must be c o n s i d e r e d i n d e c i d i n g what length of exposure flies  to use i s the p e r i o d r e q u i r e d to remove  from a b i r d between exposures.  p i e s about  10 minutes,  Since t h i s u s u a l l y  occu-  a maximum of four 30-minute exposures,  seven 15-minute exposures, or nine 10-minute exposures could be made i n the usual 3-hour c r e p u s c u l a r p e r i o d a v a i l a b l e . b a s i s of the data i n Table XIV such exposure 39, 64, and 70 s i m u l i i d s , r e s p e c t i v e l y .  On the  s e r i e s would  yield  The f o r e g o i n g compari-  sons s t r o n g l y c o n t r a i n d i c a t e the use of exposures of prolonged duration. 2. C o l l e c t i o n of C u l i c o i d e s The date presented i n Table XV  f o r c o l l e c t i o n of  C u l i c o i d e s suggest c o n c l u s i o n s s i m i l a r to those f o r s i m u l i i d s . They imply even more marked decrease i n c o l l e c t i o n with i n c r e a s e d exposure  time.  The data i n Table XV suggest that  by the end of 15- or 30-minute exposures the  efficiency  some midges had  left  grouse, and i f t h i s i s t r u e , that r e c r u i t m e n t decreased i n  prolonged exposures.  The  apparent decrease i n recruitment may  170 be  p a r t l y e x p l a i n e d by the f a c t  carried and  out before peak d a i l y  that most of the exposures were  activity  of C u l i c o i d e s occurred,  t h e r e f o r e , at a given time only a small segment of the  ambient f l i e s were f o r a g i n g . Table  XV.  C o l l e c t i o n of C u l i c o i d e s from a r b o r e a l evening exposures of d i f f e r e n t d u r a t i o n at the same s i t e i n three y e a r s . *  I U  (2) ,  I U  (2)/(l)  ( 3 ) / ( l T . ' (2)/(3)  Ye ar 1961  28  295  282  10  10  1.0  1962  16  245  122  15  7.6  2.0  1964**  21 '  630  113  30  5.4  5.6  * Based on the 5 most productive #8, Nanaimo Lakes. ** Basic data Explanation (1) (2) (3) (2) / ( l ) (3) / ( l ) (2)/(3)  D.  from Woo  (55).  F l u c t u a t i o n s of Vector  Populations,  Creek Simuliids.  August 17,  site  of columns: Number of exposures. T o t a l exposure time ( i n minutes). T o t a l number of C u l i c o i d e s captured. Average exposure d u r a t i o n ( i n minutes). Average number of C u l i c o i d e s per exposure. T h e o r e t i c a l time r e q u i r e d to capture one, C u l i c o i d e s ( i n minutes).  Seasonal 1. Trout  are  evenings during June at  1959  Exposures of grouse from May  r e v e a l e d on the b a s i s of captures,  29 to that the  major p e r i o d of abundance of g r o u s e - b i t i n g s i m u l i i d s was between J u l y 12 and served  August 13.  on the wing by June 29,  on June 30 proved to be S.  However, s i m u l i i d s were and  aureum.  s e v e r a l taken  by  ob-  sweeping  Moreover, a l l exposures  171 up to J u l y 12 were t e r r e s t r i a l , sures at stream l a t e r to be  This  proved  an u n u s u a l l y p r o d u c t i v e c o l l e c t i o n f o r t e r r e s t r i a l  nearby.  have i n d i c a t e d  abundant emergence from the  Therefore, i t i s probable  s i m u l i i d s were p l e n t i f u l late  s e r i e s of three expo-  l e v e l J u l y 6 .yielded 6 s i m u l i i d s .  exposures and may stream  and one  that g r o u s e - b i t i n g  somewhat before J u l y 12,  likely  by  June. The  onset of summer abundance of s i m u l i i d s  occurred  s h o r t l y a f t e r the extended s p r i n g f l o o d of Trout Creek had abated.  Between May  8 and May  15, when the water l e v e l began  to r i s e  n o t i c e a b l y , a few u n i d e n t i f i e d b l a c k : f l i e s were i n e v i -  dence.  During  the p e r i o d of f l o o d , from mid-May to mid-June,  b l a c k f l i e s were not seen.  A f t e r August 13,  s i m u l i i d s were  very s c a r c e , but i t i s not known whether l a t e summer occurred.  May  and  June of 1959  were abnormally  (12 cloudy days i n June, on 6 of which r a i n  resurgence  c o o l and  fell)  and  wet  the  extensive p e r i o d s of very hot, dry weather which u s u a l l y p r e v a i l i n J u l y and August, were much reduced. summers, floodwaters recede of g r o u s e - b i t i n g f l i e s duration.  sooner and  Probably  in, t y p i c a l  the p e r i o d of abundance  commences e a r l i e r and  i s of s h o r t e r  (In the e x c e p t i o n a l l y a r i d summer of 1960,  were already at a very low Culicoides. from grouse i n 1959,  l e v e l when f i r s t  between J u l y 22 and August 10. better collecting  sampled,' J u l y  B i t i n g midges were very r a r e l y  but i n 1960  techniques  a total  simuliids.  of 40 was  While t h i s may  22.)  taken  captured  or may  not  reflect  i n the second year, i t does demon-  s t r a t e that the midges continue to attack grouse,  although i n  low numbers, a f t e r black f l y p o p u l a t i o n s have dwindled  markedly.  172 2. Nanaimo Lakes Simuliids.  The most extensive s e r i e s of  i n t h i s r e g i o n encompassed the p e r i o d May E a r l y exposures stream, 11.  where s i m u l i i d s were f i r s t  11 to May  10 to August 13,  c o l l e c t e d from grouse  about 1 s i m u l i i d per exposure  24 at which time  i t was  May from  found that s i t e s i n  other l o c a t i o n s were much more p r o d u c t i v e . used  1961,  were c a r r i e d out i n a red cedar near a small  C o l l e c t i o n s averaged  May  exposures  Since the  first-  s i t e r e t a i n e d i t s low p r o d u c t i v i t y when t e s t e d p e r i o d i c a l l y  throughout  June, black f l i e s were probably more abundant  else-  where during t h i s e a r l y p e r i o d than c o l l e c t i o n s i n d i c a t e d . determined  by the number of f l i e s  captured, the major p e r i o d  of  abundance of g r o u s e - b i t i n g s i m u l i i d s extended  to  June 24  of  3-5  i n 1961,  flies  and to June 18 i n 1962,  per 10 minute  from l a t e  exposure. the dominant  s p e c i e s captured and outnumbered S. aureum by 3:1 (55) i n d i c a t e  J u l y 1, 1964).  a r a t i o . 6f 1:1  to  sustained this  2 weeks before d e c l i n i n g r a p i d l y .  June, 1961,  the  dropped to a very low l e v e l  About the end of flies  (averaging 1 f l y per 3 a r b o r e a l  exposures) which obtained throughout disappeared during t h i s  end  l e v e l f o r approxi-  p o p u l a t i o n s of adult d e n d r a g a p o p h i l i c black  interval  July.  S, aureum a l l but  (38 minus:  Between J u l y 30 and August 13, were captured  and  In the three years f o r which data have been  the f i r s t week of June and  mately  6:1.  between June 13  a c q u i r e d S. aureum reached maximum numbers at.about of  May  with an"average  Throughout t h i s p e r i o d C. minus was  (Woo's data  As  1 aureum).  1961,  (1 per 3 a r b o r e a l exposures);  only 14  flies  a l l were S. aureum.  173 This  is interpreted  this  species  occurred  on August 14  and  15,  date no data are fire  1961,  y i e l d e d no  available.  (Closure  curtailment  August 4,  days were sunny and  stimulating  1961,  hot  vanished. • Exposures  s i m u l i i d s and of the  after this  f o r e s t due  to  of f i e l d research.)  (75-90° max,); conceivably,  . • Bet-  no p r e c i p i t a t i o n ' occurred  and  besides  stages of s i m u l i i d s by abnormal diminution  the watercourses i n t h i s r e g i o n .  I f resurgence of the  S.  i n mid-summer i s an annual phenomenon I t may  prolonged and  of aureum  be more  l e s s i n t e n s i v e i n wetter summers.  Culicoides. i n the  of  emergence, t h i s weather could have r e s u l t e d i n some  l o s s of aquatic  population  a minor resurgence  a f t e r C. minus had  hazard n e c e s s i t a t e d  ween J u l y 7 and the  as i n d i c a t i n g that  Grouse-biting  Nanaimo Lakes r e g i o n  C u l i c o i d e s were present  throughout the p e r i o d mid-May to  mid-August, 1961.  They were most abundant i n exposures made  between May  June 28 and  rately  low  31  and  l e v e l which was  Throughout the  crepuscularis group was  maintained throughout the  peak p e r i o d the  plain yellowish  species  group was  decreased t h e r e a f t e r to a mode-  predominant species  r e f e r r e d to as C u l i c o i d e s slightly  During the  p o r t i o n of C. u n i c o l o r group i n c r e a s e d , populations  was  and  the  sp.. 1.  l e s s abundant, and  encountered i n f r e q u e n t l y .  summer.  C.  low  C.  unicolor  summer the  i n the  small,  pro-  August  t h i s was- the most numerous form.  * Baranov (9) found summer outbreaks of S. columbaczense r e l a t e d to low r a i n f a l l , high a i r temperatures and low, warm water i n the Danube; p o s s i b l y , S. aureum resurgence occurs only i n hot, dry summers.  174 E.  Feeding A c t i v i t y of V e c t o r s 1. R e l a t i o n of a c t i v i t y to s t r u c t u r a l f a c t o r s and h a b i t a t Simuliids.  ,In O n t a r i o , Bennett  s p e c i e s of o r n i t h o p h i l i c black f l i e s quebecense.  ( l l ) ' found that some  such as S. l a t i p e s . S.  and S. r u q q l e s i feed p r i m a r i l y on hosts at ground  l e v e l , while others such as S. aureum. S. c r o x t o n i . S. decemarticulatum. and C. o r n i t h o p h i l i a  attack b i r d s well-above the  ground, i n t r e e s . In  the present study, the w r i t e r c a r r i e d out a r b o r e a l  and t e r r e s t r i a l  exposures i n both r e s e a r c h areas.  In 1961  when most of the t e r r e s t r i a l exposures were made, the f o l l o w i n g r e s u l t s were o b t a i n e d , May 10 - August sures produced 648 s i m u l i i d s and 223 t e r r e s t r i a l per  13:  (1.6 f l i e s  397 a r b o r e a l expo-  per 14-minute exposure),  exposures produced 60 s i m u l i i d s  (0.27 f l i e s  14-minute exposure) with the same s p e c i e s composition. -By  using 2 grouse i n simultaneous exposures, the numbers of b i t i n g flies  at ground  l e v e l and about 40 f e e t  same s i t e were compared on 5 evenings.  above ground at the These data are given i n  Table XVI. •Table XVI.  Date  ill  R e s u l t s of c o l l e c t i n g b i t i n g . f l i e s from two grouse i n simultaneous a r b o r e a l and t e r r e s t r i a l evening exposures of same d u r a t i o n .  121  Simuliids Tree Ground  12/7/59  2  20  14  1  4/6/61  3  45  17  1 .'  18/6/61  5  50  35  19/6/61  3  35  10/6/62  1  15  Culicoides Tree Ground 0  0  24  2  1  2  0  30  3  77  20  7  0  0  0  E x p l a n a t i o n of columns: (1) Number of exposures at each (2) T o t a l exposure time at each  level. level.  175 It  must  be concluded that both .S. aureum and  C. minus feed, f o r the most p a r t , w e l l above the ground. Throughout firs  the study, grouse were exposed  almost e x c l u s i v e l y ,  lower l i v i n g branches.  i n mature Douglas  and were r a i s e d to the l e v e l However, on Vancouver  of the  I s l a n d a few  exposures were c a r r i e d out i n hemlocks  and r e d cedars.  The  l a t t e r t r e e s were at the edge of dense  alder swales w i t h i n  50 feet of small watercourses i n r u f f e d grouse, r a t h e r than blue grouse h a b i t a t , and they y i e l d e d few f l i e s . exposed  frequently i n 2 t a l l  Douglas  firs  Grouse were  150 and 300 feet  from one of the cedars and numerous s i m u l i i d s were These in  f i r s were on s l i g h t l y  a Douglas  higher, considerably d r i e r  ground  f i r - w e s t e r n h e m l o c k - s a l a l a s s o c i a t i o n , and the  grouse were r a i s e d cedars.  collected.  15-20 feet higher i n the f i r s  It i s difficult  than i n the  to decide which of these v a r i a b l e s  c o n t r i b u t e d t o the g r e a t e r abundance of black f l i e s and l i k e l y a number of f a c t o r s were i n v o l v e d . o b s e r v a t i o n s suggest that a c c e s s i b i l i t y  i n the f i r s  The f o l l o w i n g  of i n d i v i d u a l  trees  may be among the more important. Most p r o d u c t i v e of f l i e s  was one Douglas  f i r , expo-  sure s i t e #8, s i t u a t e d on the p e r i p h e r y of a spur of Douglas f i r - h e m l o c k f o r e s t where i t terminated a b r u p t l y at the burntover, open, grouse range.  In 1961, p r e l i m i n a r y exposures of  grouse i n s e v e r a l other Douglas 200 yards southwest few s i m u l i i d s .  firs  i n this v i c i n i t y ,  about  of the North Nanaimo R i v e r , y i e l d e d very  In an attempt to determine whether only  #8 was frequented c o n s i s t e n t l y by large numbers of black two grouse were exposed  site flies  s i m u l t a n e o u s l y , at s i t e #8 and i n one  176 of the other 4 t a l l trees  (#9, #10,  Douglas f i r s  #11,  selected  f o r comparison.  These  and #12)  were chosen because they were  approximately the same height  as #8 and because t h e i r lower  branches, to which the grouse were r a i s e d , were about the same d i s t a n c e from the ground. From the r i v e r a d i r t road runs southward open range and b i s e c t s  the f o r e s t  spur.  across the  On the east side  of the  road the spur extends about 50 feet nearer the r i v e r than on the west the  side.  Douglas  northern l i m i t  f i r #8 is. on the west of t h i s side  from the open range. the  spur.  feet  F i r #10  of the road at  of the spur, and i s separated  from the road by a narrow c l e a r i n g ; of the road but about 50 feet  side  f i r #12  i s on the same side  southwest of #8 and about 50  feet  The other 3 s i t e s are i n the east part  i s about 25 feet  from the road and about  100  south of the open range, and surrounded' by•other c o n i f e r s ;  f i r #11  i s about 150  feet east of the road, about 50 feet south  of the open range, and i s surrounded by other c o n i f e r s ; #9 i s at the east r o a d s i d e , about 500  feet  at #8 -with the number c o l l e c t e d  and f i r  south of #8.  Table XVII compares the number of "simuliids at the other s i t e s .  collected  Exposures  at each p a i r of s i t e s were the same i n number and d u r a t i o n , were c a r r i e d out s i m u l t a n e o u s l y with 2 grouse'which were a l t e r n a t e d show that  between s i t e s .  These comparisons  o r n i t h o p h i l i c black f l i e s  around f i r #8 i n f a r g r e a t e r numbers than at nearby s i t e s . ation  of  The w r i t e r  f o r t h i s phenomenon.  and  usually  (p. 177)  were c o n s i s t e n t l y present (with only one exception)  suggests the f o l l o w i n g  l ) The p r e v a i l i n g  explan-  breezes, which  blow up the v a l l e y from the southeast, probably eddy i n t o the  177 clearing  c r e a t e d by the road ( i n the lee of the grove which  p r o j e c t s s l i g h t l y i n t o the open range on the' east side of the road).  Most s i m u l i i d s c a r r i e d by the breeze Were dropped i n  the t r e e s on the west side of the road (#8) but' some came to rest ing.  on the east side The f i l t e r i n g  (#10) as the flow c i r c u l a t e d i n the c l e a r -  effect  of the t r e e s at the edge of the road  and the edge of the f o r e s t prevented many f l i e s t r e e s deeper i n the f o r e s t road  (#9).  from r e a c h i n g  (#11, #12) and f a r t h e r along the  2) Since there are no streams w i t h i n t h i s part of  the f o r e s t the s i m u l i i d s probably had emerged from the North Nanaimo R i v e r , whence they moved upward across the open range to the f o r e s t p e r i p h e r y e i t h e r on convection c u r r e n t s or through an innate tendency to seek higher ground or a r b o r e a l  resting  places. Table XVII.  Comparison of s i m u l i i d c o l l e c t i o n s at the most p r o d u c t i v e s i t e and four other s i t e s i n i t s v i c i n i t y . Number of exposures  Simuliids collected  Date  Time  20/6/61  Dawn  3 each  20/6/61  Dusk  3 each  #8- 41; . ..#9-0 #8- 15.5 .#10--21  21/6/61  Dawn  3 each  #8- 26; . #10--2  22/6/61  Dusk  3 each  #8- 20; #10--1  22/6/61  Dusk  3 each  #8- 31; #11--6  24/6/61  Dusk  2 each  .#8 -9; #11--3  24/6/61  Dusk  2 each  #8 -7; #12--1  Culicoides.  Bennett  (ll)  noted that s e v e r a l  species  of C u l i c o i d e s were much more abundant i n a r b o r e a l s i t e s than at ground l e v e l .  As i n d i c a t e d by Table XVI (p. 174) s i m i l a r  results  178 were obtained i n the present study, although these and  other  data r e v e a l l e s s discrepancy i n abundance of C u l i c o i d e s at ground l e v e l and t r e e l e v e l than f o r s i m u l i i d s .  During  June,  when C u l i c o i d e s are most abundant, twice as many were captured, per exposure, terrestrial  i n t r e e s as on the the ground.  c o l l e c t i o n s were almost  In other months  negligible.  Since the  midges from both e l e v a t i o n s were preserved together the s p e c i e s composition  of ground-feeders  2. R e l a t i o n of a c t i v i t y  i s unknown.  to environmental  factors  V a r i o u s s t u d i e s have shown that the feeding behavior of  black f l i e s v a r i e s i n i n t e n s i t y r e l a t i v e to changes i n i l l u m -  i n a t i o n , temperature, and wind v e l o c i t y . interdependent,  r e l a t i v e humidity,  atmospheric  Most of .these f a c t o r s are i n t e r r e l a t e d  and t h e r e f o r e i t i s very d i f f i c u l t  t h e i r i n d i v i d u a l e f f e c t s on s i m u l i i d behavior. s p e c i e s , the same f a c t o r or combination evoke d i f f e r e n t responses  (23).  feeds mainly during d a y l i g h t eratures  decreased  uliids  to assess  From s p e c i e s to  of f a c t o r s may  For i n s t a n c e , S.  even  arcticum  and' i s most numerous- a't high temp-  during the day  i l l u m i n a t i o n and temperature  and i s most a c t i v e i n the of e a r l y evening  (20).  ( l l ) c o l l e c t e d woodland s p e c i e s of o r n i t h o p h i l i c i n g r e a t e s t numbers during l a t e evening,  Peterson ing  or  (46), whereas S. venustum i s a c t i v e i n the shade at  moderate temperatures  Bennett  pressure,  (54)  activity  Wolfe and  suggested  and Wolfe  simand  that the d i u r n a l rhythm of s i m u l i i d  i s d i r e c t l y r e l a t e d to changing  light  feed-  intensity.  Peterson d i s c o v e r e d also that black f l i e s move upward  toward the top of the f o r e s t cover i n the evening c l o s e r to ground l e v e l  i n the e a r l y morning.  and  descend  This behavior  may  179 have c o n s i d e r a b l e  s i g n i f i c a n c e i n the transmission of avian  filarioids. The  g e n e r a l l y c r e p u s c u l a r h a b i t s of a n t h r o p o p h i l i c  C u l i c o i d e s are w e l l known, though c o n s i d e r a b l e p r e f e r r e d feeding times occurs: the n i g h t , while and  latitude i n  C. m i l n e i a t t a c k s man during  C. qrahami b i t e s i n e a r l y morning and-evening  i n d u l l , wet d a y l i g h t hours (41).  species c o l l e c t e d by Bennett  The 6 o r n i t h o p h i l i c  ( l l ) using evening  were most abundant a f t e r 2000 hours  (sunset  exposures only,  2045-2100), and  3 species were more a c t i v e between 2200 hours and midnight. In the present  study,  an attempt was made to d e t e r -  mine whether the feeding a c t i v i t y of g r o u s e - b i t i n g s i m u l i i d s and  C u l i c o i d e s during  the major p e r i o d of t h e i r abundance  showed any r e l a t i o n s h i p to 2) temperature, and  l ) time of day and i l l u m i n a t i o n ,  3) wind.  Time of day.and i l l u m i n a t i o n .  Exposures made i n the  Nanaimo Lakes r e g i o n at i n t e r v a l s from 0900 to 1500 hours PST y i e l d e d 4 s i m u l i i d s on a calm sunny' day (maximum temperature 78°F), day  and 7 s i m u l i i d s and 2 C u l i c o i d e s on a calm  (max. temp. 72°F).  In c o n t r a s t , evening  same days produced numerous f l i e s .  overcast  exposures on the  Apparently  then,  none of  the g r o u s e - b i t i n g black f l i e s or midges engages i n i n t e n s i v e feeding  a c t i v i t i e s between mid-morning and mid-afternoon, but  perhaps a c t i v i t y this  i n c r e a s e s s l i g h t l y i n cloudy weather during  period. In June, i n the North Nanaimo R i v e r v a l l e y ,  occurs 1900  l a t e , at about 0600 hours, and sunset  sunrise  e a r l y , at about  hours PST, due to the presence of Blackjack Ridge on the  180 east and Mt.  de Cosmos on the west.  mid-afternoon u n t i l n i g h t f a l l  Exposures  (at 2100  to 2130  r e v e a l e d that black f l y f e e d i n g a c t i v i t y about  1630  hours.  on, with the m a j o r i t y b i t i n g  In the next hour a c t i v i t y  s i m u l i i d s were recovered a f t e r  conducted' from hours),  i n c r e a s e d from from 1730  to 2030  decreased, and only a few  dark.  C u l i c o i d e s . on the other hand, commenced feeding later  i n the evening.  e a r l y as 1730 1845  a few were taken from grouse  hours, the time of f i r s t  activity  Culicoides  observed to commence f e e d i n g a c t i v i t y one-half to  one hour e a r l i e r Ten hours,  Peak  and 2200 hours, but feeding continued  l e a s t u n t i l midnight with reduced i n t e n s i t y ;  sp. 1 was  than C. c r e p u s c u l a r i s  10-minute exposures  (when f e e d i n g a d t i v i t y  group.  between 0550 and 0900  virtually  c l e a r mornings June 20-24, 1961  produced  ceased) 86  (S.  aureum and C. minus) and 4 C u l i c o i d e s .  ten  10-minute exposures  between 1730  on  still,  simuliids By  comparison,  and 2200 hours on  c l e a r evenings i n the same p e r i o d y i e l d e d 97 s i m u l i i d s 170 C u l i c o i d e s .  In these and other e a r l y morning  the l a s t b l a c k f l y was  The  still, and  exposures  captured at the f o l l o w i n g times:  0800, 0830, 0840 and 0845  Co  as  capture ranged between  and 2000 hours PST f o r 12 n i g h t s i n June,  occurred between 1900 at  While  0740,  PST.  f o r e g o i n g o b s e r v a t i o n s show that S_. aureum and  minus e x h i b i t two p e r i o d s of intense feeding a c t i v i t y ,  e a r l y p e r i o d extending to mid-morning, the l a t e  period  n i g h t f a l l , and. suggest that i n the i n t e r v e n i n g daytime of quiescence  l i m i t e d a c t i v i t y may  resume under reduced  the  until period  181 illumination.  C u l i c o i d e s are a p p a r e n t l y r e s t r i c t e d to one  major p e r i o d of a c t i v i t y commencing at dusk and extending f o r an undetermined  time i n t o the n i g h t .  Temperature.  At the t i m e s : l i s t e d f o r l a s t  morning  capture of s i m u l i i d s , the a i r temperatures: i n order, were 58, 66, 70, 62, and 75°F.  In evening exposures s i m u l i i d s were  taken at temperatures as high as 80°F, but no h i g h e r temperatures were r e c o r d e d at negative exposures, and few s i m u l i i d s were captured w i t h i n t h i s range of temperatures at other times of the day.  These o b s e r v a t i o n s suggest that  diurnal  a c t i v i t y of these species may be independent of temperature (over a c o n s i d e r a b l e range, at l e a s t ) , the  c o n t e n t i o n of Wolfe  and tend t o support  and Peterson (54), that a c t i v i t y i s  c o n t r o l l e d mainly by g r a d u a l l y changing l i g h t Wind. are of  intensity.  In the Trout Creek v a l l e y summer evenings  u s u a l l y accompanied  by moderate to strong breezes.  the few calm evenings when b l a c k f l i e s were  y i e l d e d about 200 s i m u l i i d s  One  abundant  (mostly S. aureum) i n 2 expos-  ures; numbers taken when wind v e l o c i t y was about 20 m.p.h. (estimated) were of c o n s i d e r a b l y lower magnitude, winds over 30 m.p.h. about and no f l i e s  and i n  (est.) the f i r branches were b u f f e t t e d  were captured.  On the North Nanaimo R i v e r grouse range,  strong  winds seldom occur during the summer, and many evenings are calm.  No a p p r e c i a b l e d i f f e r e n c e i n the number of s i m u l i i d s  captured was noted between calm and windy evenings. In June  the same area, 12 exposure-evenings between  14 and June  27, 1961 and 1962 (combined)  y i e l d e d an  182 average  of 30 C u l i c o i d e s per evening when the weather was  clear  and calm, but b c l e a r , windy evenings and 1 o v e r c a s t windy evening i n t h i s p e r i o d produced  a t o t a l of only b C u l i c o i d e s .  From these o b s e r v a t i o n s i t i s concluded that the feeding a c t i v i t i e s of C u l i c o i d e s are suppressed by winds of c o n s i d e r ably lower v e l o c i t y than those which reduce  simuliid  activity.  183 VI. The  EPIZOOTIOLOGY  d i v e r s e f a c t s and c o n c l u s i o n s concerning the  b i o l o g y of the f i l a r i o i d s ,  t h e i r t e t r a o n i d hosts and t h e i r  d i p t e r a n v e c t o r s which were d i s c u s s e d independently i n previous  s e c t i o n s are here considered  i n r e l a t i o n t o each  other.  Since much of the f i e l d r e s e a r c h i n v o l v e d Micro-  filaria  sp. B, blue grouse, and s i m u l i i d s on Vancouver I s l a n d ,  this  complex w i l l be considered i n g r e a t e s t d e t a i l ;  filarioids, picture  other  hosts, and l o c a l i t i e s w i l l be brought i n t o the  f o r comparison.  A., Blue  Grouse on Vancouver I s l a n d When these  grouse descend i n s p r i n g from the winter  range to t h e i r breeding  areas on the d e f o r e s t e d f l a t l a n d s and  lower s l o p e s , almost a l l the a d u l t s and y e a r l i n g s harbor e i t h e r S k r i abinocta f l e x i v a q i n a l i s or M i c r o f i l a r i a (?=Splendidofilaria  sp. B  s p . ) ; most adult males, but few year-  l i n g s c a r r y double i n f e c t i o n s .  During  the p e r i o d when  t e r r i t o r i e s become e s t a b l i s h e d and n e s t i n g s i t e s are s e l e c t e d i t i s u n l i k e l y that f i l a r i a l  transmission  occurs,  because few o r n i t h o p h i l i c d i p t e r a n s are emerging and a i r temperatures over extended periods may be below the minimum necessary  f o r development of f i l a r i a l Microfilaria  biting  flies  sp. B.  larvae.  By l a t e May, p o p u l a t i o n s of  are i n c r e a s i n g , but the g r o u s e - b i t i n g s i m u l i i d s  comprise Cnephia minus almost e n t i r e l y . few  larvae of Mf. sp. B can develop  Since  comparatively  i n an i n f e c t e d C. minus.  and because daytime temperatures are s t i l l  moderately low  (therefore., l a r v a l development r e q u i r e s s e v e r a l weeks), t r a n s m i s s i o n i s probably  n e g l i g i b l e before  e a r l y June.  In e a r l y June, a i r temperatures r i s e , t i o n s of C. minus and  and  S. aureum i n c r e a s e markedly, thus  g r e a t l y enhancing the t r a n s m i s s i o n p o t e n t i a l of Mf, The  next 2 to 3 weeks are probably  transmission  y e a r l i n g males.  the host  population,  contact i s i n c r e a s e d , but  t a n t l y , the p r o b a b i l i t y of v e c t o r - u n i n f e c t e d host is greatly increased.  B.  sp. B are a t t r a c t e d to young  c h i c k s , then when the chicks enlarge p r o b a b i l i t y of v e c t o r - h o s t  sp.  the most f a v o r a b l e f o r  of t h i s species to a d u l t and  I f the v e c t o r s of Mf.  filarial  popula-  I t appears that the  the  concomi-  contact  success  of  t r a n s m i s s i o n to newly hatched grouse chicks  should  be i n f l u e n c e d s t r o n g l y by the degree of synchrony between peak abundance of v e c t o r s and peak hatching us c o n s i d e r two  types  c h i c k s appear before this  case,  as, f l i e s  of synchrony.  of c h i c k s .  Type 1: most of the  the v e c t o r populations  enlarge.  The  on  r a t i o of i n f e c t e d to u n i n f e c t e d  v e c t o r s w i l l be moderately low. few  In  become abundant many newly emerged  i n d i v i d u a l s of the v e c t o r species w i l l be feeding u n i n f e c t e d grouse.  Let  Therefore,  correspondingly  c h i c k s w i l l become i n f e c t e d e a r l y . Type 2: most of the c h i c k s hatch  t o r s have been abundant f o r s e v e r a l weeks.  after' the The  r a t i o of  i n f e c t e d to u n i n f e c t e d v e c t o r s w i l l be c o n s i d e r a b l y than i n the previous  case,  and  p r o p o r t i o n of the c h i c k s should  vec-  higher  a correspondingly l a r g e r acquire i n f e c t i o n s e a r l y .  185 At f i r s t  glance, the r e l a t i o n s h i p of v e c t o r to  young c h i c k i n the t r a n s m i s s i o n of Mf. Type 2 synchrony.  sp. B seems'to  C. minus and S. aureum have been  be of abundant  f o r upwards of a week before the c h i c k s hatch, and have had s u f f i c i e n t time to feed on i n f e c t e d adult or y e a r l i n g L a r v a l development  i n s i m u l i i d s probably r e q u i r e s  grouse.  10-14  days at the temperatures of e a r l y to mid-June; t h e r e f o r e , one would  expect i n f e c t i v e  inoculum to be present in., the  b l a c k f l i e s when most of the chicks are newly  hatched.  Many chicks should become i n f e c t e d w i t h i n a few days of h a t c h i n g ; i n a c t u a l i t y , t h i s i s not the case.' None of the large number of c h i c k s removed a l i v e between June captivity;  from the grouse range  11 and J u l y 1 developed m i c r o f i l a r a e m i a s i n  o b v i o u s l y they were captured before they had  been b i t t e n by i n f e c t e d v e c t o r s .  ' •  The f o l l o w i n g r e l a t e d aspects of the h a b i t s of c h i c k s and of v e c t o r s probably r e s u l t c h i c k contact during June:  i n minimal v e c t o r -  l ) U n t i l they are about 2 weeks  o l d , c h i c k s are i n c a p a b l e of e f f i c i e n t restrictedly. terrestrial; much l e s s abundant  simuliid  and are .  but S. aureum and C. minus are  at ground  l e v e l than i n t r e e s . .  a c t i v i t y commences each day when d i r e c t the dew  flight  and the a i r temperature r i s e s ;  2) Chick  s u n l i g h t evaporates but by t h i s  time  a c t i v i t y has d e c l i n e d from i t s early-morning peak. The presence of Leucocvtozoon i n a few c h i c k s i n  June  almost undoubtedly i n d i c a t e s that, some simuli'id-chick  contact occurs before the grouse are one month o l d , perhaps much of t h i s as a r e s u l t  of egg-laden female  simuliids  186 passing  low over grouse range to o v i p o s i t i n the r i v e r s .  Probably  some v e c t o r - c h i c k contacts i n t h i s p e r i o d  place i n the evenings, and  black f l i e s  when temperatures are s t i l l  are a c t i v e .  the most favourable  M i l d , overcast  take elevated  days may  conditions f o r transmission,  provide  since  s i m u l i i d s tend t o drop near the ground during the day and become a c t i v e i n cloudy  weather.  Since C. minus i s c o n t i n u o u s l y as a low p o p u l a t i o n , is  present,  throughout most of J u l y , t h i s  though species  l i k e l y r e s p o n s i b l e f o r mid-summer t r a n s m i s s i o n to the  r a p i d l y growing c h i c k s . In e a r l y August a resurgence of S. aureum occurs. Despite  i t s low d e n s i t y , there  believing  that t h i s p o p u l a t i o n  able t r a n s m i s s i o n efficient  are s e v e r a l reasons f o r i s responsible f o r consider-  of Mf. sp. B.  intermediate  host  l ) S. aureum i s a h i g h l y  and i s capable  large numbers of larvae per f l y . high; development t o i n f e c t i v e week.  3) Chicks  wander widely,  of supporting  2) A i r temperatures are  stage  r e q u i r e s l e s s than a  are a c t i v e e a r l y and l a t e i n the day, they  and they  occasionally f l y into trees.  Some adult male blue grouse are present  on the  range when the resurgence of S. aureum begins;-however, by mid-August v i r t u a l l y  a l l have departed.  This means that  e a r l y i n August the newly emerged black f l i e s acquire  probably  inoculum from cocks as w e l l as hens, but l a t e r ,  only the c h i c k s and hens are a v a i l a b l e to r e c e i v e larvae.  infective  187 The  higher  prevalence of Mf.  sp. B (Table  adult males than i n adult females or i n y e a r l i n g s which could have acquired is  most e a s i l y  t h e i r i n f e c t i o n s only  a t t r i b u t e d to the r e l a t i v e  these groups to i n f e c t e d v e c t o r s . are  the most f r e q u e n t l y  of t h e i r d a i l y  attacked  accessible on the  the  area  contiguous, "hooting"  occupied  and  bears few  chicks)  availability  are  of  abundant i s  In the high grouse  a s s o c i a t e d with s l i g h t e l e v a t i o n s and  as  s i t e s i n which they are r e a d i l y  to these v e c t o r s .  are  (most of  by s i m u l i i d s 'because much  open "burns", male t e r r i t o r i e s  frequently  in  Presumably,- adult males  a c t i v i t y while vectors  r e s t r i c t e d to "hooting"  IX)  populations  are r a t h e r small s i t e s are  and  and  often  young Douglas  firs,  by grouse i s r e l a t i v e l y homogeneous  obstructions  to the movement of  vectors.  Y e a r l i n g males on the breeding range are u s u a l l y found wandering through the t e r r i t o r i e s they l i k e l y  of the  cocks; as a r e s u l t  encounter pockets of i n f e c t e d vectors The  lower prevalence of Mf.  frequently.  sp. B i n adult hens  than i n adult  cocks may  p e r i o d during  most of which the hens are r e s t r i c t e d  their terrestrial  be r e l a t e d to the  nests.  On  the other  s e v e r a l s t u d i e s i n d i c a t e that the of f i l a r i a l  i n f e c t i o n s recorded  f a c t o r s besides  lengthy  incubation  hand, r e s u l t s of  differential  here may  prevalence  reflect  differential availability.  other  Dalmat  (19)  showed that i n t e n s i t y of s i m u l i i d - f e e d i n g on humans r e l a t e d to the and Wolfe and  amount of l i g h t r e f l e c t e d from the Peterson  (54)  to  was  skin,  found that i n the morning  and  evening l i g h t - s k i n n e d people were b i t t e n more f r e q u e n t l y  188  than those with dark c o l o u r a t i o n , whereas during l a t t e r i n d i v i d u a l s were attacked Fallis  and Smith (24) provided  ornithophilic  more o f t e n .  convincing  the day the  Recently,  evidence that some  s i m u l i i d s are s t r o n g l y a t t r a c t e d - t o  hosts by chemical o l f a c t o r y s t i m u l i .  their  I t might be r a t h e r  p r o f i t a b l e i n future to i n v e s t i g a t e whether the d i f f e r e n c e s i n pigmentation, s i z e , and p h y s i o l o g i c a l s t a t e which e x i s t i  among cocks, hens and chicks r e s u l t i n d i f f e r e n t " a t t r a c t i o n p o t e n t i a l s " f o r s i m u l i i d s , and i f so, t o what extent i n f l u e n c e the t r a n s m i s s i o n S k r i abinocta  of Mf. sp. B.  flexivaqinalis.  u n i c o l o r group i s present  Since  Culicoides  on the grouse range' from e a r l y  June u n t i l mid-August at l e a s t , t r a n s m i s s i o n probably occurs throughout t h e i r f i r s t the  same f a c t o r s which i n f l u e n c e  sp.  B no doubt a f f e c t t r a n s m i s s i o n The  these  to chicks  summer.  Many of  the t r a n s m i s s i o n of t h i s  of Mf.  species.  data i n Table IX (p, 114) i n d i c a t e that i n  a d u l t s of e i t h e r sex and i n both y e a r l i n g s and a d u l t s , Mf.  sp.' B i n f e c t i o n s s l i g h t l y outnumber Sk. f l e x i v a q i n a l i s  infections.  For the f o l l o w i n g reasons one might expect'  t h i s to be the case: population  l ) the moderately small  f o r Sk,. f l e x i v a q i n a l i s . 2) the use by C u l i c o i d e s  of one, r a t h e r than two feeding greater  periods  e f f e c t of wind i n suppressing  4) the i n a b i l i t y filariae  vector  d a i l y , 3) the  Culicoides  of C u l i c o i d e s to ingest  activity,  as many micro-  per i n d i v i d u a l , and 5) the longer  developmental  p e r i o d of Sk. f l e x i v a q i n a l i s i n i t s intermediate  host.  189 However, the d i f f e r e n c e s i n prevalence two  filarioids  are not s t a t i s t i c a l l y  of these  significant.  I t may be  that the f o l l o w i n g aspects of the b i o l o g y of C u l i c o i d e s m i t i g a t e some of the negative i n f l u e n c e s by f a v o u r i n g vector-host contact: in  l ) the tendency f o r C u l i c o i d e s t o feed  moderate numbers at g r o u n d - l e v e l where most of the  activities  of the host take p l a c e ,  feeding a c t i v i t i e s  2) p r o l o n g a t i o n of the  of C u l i c o i d e s a f t e r n i g h t f a l l , when the  grouse are r o o s t i n g on the ground or i n the young Douglas firs,  motionless,  3) continuous  and t h e r e f o r e e a s i l y attacked, and  presence  of the v e c t o r p o p u l a t i o n during the  summer. B.  Ruffed Grouse on Vancouver I s l a n d The  f r e q u e n t l y a r b o r e a l h a b i t s of r u f f e d  would seem t o predispose few  them to f i l a r i a l  i n f e c t e d b i r d s were found.  t y p i c a l of f i l a r i a l mission f a i l u r e effect  prevalence  i s probably  infections; yet,  I f the sample' examined was i n ruffed  grouse,"trans-  a f u n c t i o n of the i s o l a t i n g  of t h e i r h a b i t a t , t h e i r low p o p u l a t i o n d e n s i t y , and  the e x i s t e n c e of Type 1 v e c t o r - c h i c k synchrony. contact i s probably  flies  Vector-host  l e s s frequent i n the dense [aider swales  which r u f f e d grouse occupy than on the a d j o i n i n g and  grouse  c a r r y i n g blue grouse inoculum  "burns",  may be f i l t e r e d out  near the p e r i p h e r y . In i n t e r p r e t i n g  the e p i z o o t i o l o g y of the f i l a r i -  oids of blue grouse the p o s s i b i l i t y  of e a r l y autumn t r a n s -  m i s s i o n on or near winter range was not d i s c u s s e d . is  There  a dearth of knowledge of the h a b i t s of the m i g r a t i n g  190 grouse, the existence  of s u i t a b l e v e c t o r s , the environmental  c o n d i t i o n s and even the l o c a t i o n of the winter t h e r e f o r e , at present mission  i t i s not worth s p e c u l a t i n g on t r a n s -  at high e l e v a t i o n s .  transmission  ranges;  Should i t be found t h a t some  does occur w e l l above the summer range,  this  would f u r t h e r e x p l a i n why i n f e c t i o n s are more p r e v a l e n t i n blue grouse than i n r u f f e d . C.  Tetraonids  i n the Okanagan Region  U n t i l the s i t e s of the adults of M i c r o f i l a r i a sp, B i n i t s hosts  are known i t cannot be d e f i n i t e l y  determined whether t h i s the p r o v i n c e .  species occurs  Whether there  i n the i n t e r i o r of  are 2 or 3 species of f i l a r i -  oids i n the t e t r a o n i d s of t h i s r e g i o n does not a l t e r the fact Sk.  that none approaches the prevalence  of Mf-, sp. B or  f l e x i v a q i n a l i s i n Vancouver I s l a n d blue grouse.  filarioids  i n general  seem to be l e s s s u c c e s s f u l i n the  i n t e r i o r of the province  than on Vancouver I s l a n d .  S e v e r a l c o n d i t i o n s are a c t u a l l y more for  Avian  the t r a n s m i s s i o n of grouse f i l a r i o i d s  favourable  i n regions  such  as the Trout Creek V a l l e y than on the Vancouver I s l a n d "burns".  In the f i r s t  p l a c e , one s i m u l i i d , S. aureum.  which i s a h i g h l y e f f i c i e n t v e c t o r f o r Mf. sp. B on Vancouver I s l a n d , i s c o n s i d e r a b l y more abundant at Trout Creek.  Secondly, on the semi-open, f o r e s t e d benchlands,  blue grouse are much more a r b o r e a l i n h a b i t s .  Therefore,  for  should be  any i n d i v i d u a l grouse, vec t o r - h o s t  much more  frequent.  contact  191 On the other hand, a complex of the f o l l o w i n g f a c t o r s operates region:  to depress  l ) Grouse of any  d e n s i t y than  transmission e f f i c i e n c y  s p e c i e s are present  i n much lower  are the blue grouse on Vancouver I s l a n d "burns".  2) S i m u l i i d s do not become abundant u n t i l the s p r i n g f l o o d has developed  and  subsided.  By t h i s time the c h i c k s are w e l l  f o r these  flies  frequent winds, high temperatures and  nature  of t h i s r e g i o n operate  s u r v i v a l and breeding  h a b i t s of C u l i c o i d e s .  grouse examined was carry simuliids  and  the r a t h e r  Consequently,  i n f e c t i o n s of  that none of the s h a r p - t a i l e d  infected.  4)  Thermal u p d r a f t s tend to  from t h e i r breeding  streams to higher  grouse range at 3000 feet near Oyama where the p o s s i b l e source  of these  f l i e s was  about 1500  lower).  Yet,  The  for  as summer progresses  known.  abundance of t h i s  found  vegetation.  be that S. aureum transmits t h i s  the time of appearance of f o u r t h - s t a g e  The  blue  On the b a s i s of c i r c u m s t a n t i a l  Creek grouse c h i c k s c o i n c i d e s extremely of  nearest  i d e n t i t y of the v e c t o r s f o r S p i e n d i d o f i l a r i a  p e c t o r a l i s i s not i t may  on  2 to 4 miles d i s t a n t and  grouse move downwards i n search of greener  evidence  Sk.  ( a n t h r o p o p h i l i c black f l i e s were encountered  feet  arid  against the feeding h a b i t s ,  i s not s u r p r i s i n g to f i n d that few  f l e x i v a q i n a l i s occur,  already  - Type 1 synchrony.  3) The  elevations  prolonged  t h e i r presence has markedly d i l u t e d the  low d e n s i t y of inoculum  it  in this  filarioid,  larvae i n Trout  w e l l with the p e r i o d  simuliid.  s i n g l e r u f f e d grouse i n which the w r i t e r  S. p e c t o r a l i s was  collected  at moderately high  192 elevation.  I t i s probable that the r u f f e d grouse  i n the  i n t e r i o r of B. C. harbor few i n f e c t i o n s with S.' p e c t o r a l i s because  most r e s i d e i n the r i p a r i a n growth and thereby are  rather e f f e c t i v e l y attack blue  isolated  from contact with f l i e s  grouse. It  is difficult  to account  f o r the absence of  S. p e c t o r a l i s from c o a s t a l B r i t i s h Columbia. appears, Mf. sp. B ( ? = S p l e n d i d o f i l a r i a of  that  I f , as i t  sp.) occurs i n grouse  the i n t e r i o r as w e l l as on the coast, then i t could be  argued that S. p e c t o r a l i s i s absent from the coast i t s v e c t o r s do not occur t h e r e .  because  Obviously, the d i s t r i b u -  t i o n a l r e s t r i c t i o n of S. p e c t o r a l i s cannot be e x p l a i n e d at present. D,  Non-Tetraonid  Birds  The p h a s i a n i d s examined from B r i t i s h did  not harbor f i l a r i o i d s ,  Columbia  but the sample s i z e f o r each  s p e c i e s was not large enough to warrant  c o n c l u d i n g that they  never acquire the f i l a r i a s e s of t e t r a o n i d s . Apparently, non-gallinaceous b i r d s ' i n B r i t i s h Columbia alis .  are p a r a s i t i z e d by n e i t h e r Mf. sp. B nor S. p e c t o r -  I f each form of m i c r o f i l a r i a  found i n the non-  g a l l i n a c e o u s b i r d s r e p r e s e n t s one s p e c i e s of f i l a r i o i d , then some show r a t h e r broad host t o l e r a n c e (e.g., Form 5 i n Tyrannidae, Turdidae, Thraupidae, and F r i n g i l l i d a e ) , and yet  none p a r a s i t i z e s the Tetraonidae.  Simulium  Bennett  ( l l ) collected  aureum from 14 s p e c i e s of b i r d s of 10 f a m i l i e s i n  O n t a r i o ; t h e r e f o r e , i t i s u n l i k e l y that the absence of  193 Mf.  sp. B from n o n - t e t r a o n i d  any  host  b i r d s were c o l l e c t e d i n the  of i n f e c t e d grouse, probably Mf.  S. p e c t o r a l i s are r e s t r i c t e d physiological  to the  described  Tetraonidae alone  ( i t was  originally  Moreover, i f M i c r o f i l a r i a e x h i b i t s broad  I t s absence from most of the  ceous b i r d s examined may  seem to be  sp. B a'nd  Tetraonidae by a  f l e x i v a q i n a l i s . t h i s species  tolerance.  preferences  have been due  of C u l i c o i d e s . and attacked  2) the  to  l) f e e d i n g  fact  that small  l a r g e r number of species  (ll).  Columbia comprise a  are much more numerous on V a n c o u v e r I s l a n d  i n t e r i o r of the province.  Since  filarial  b i r d s are c o n s i d e r a b l y  more  on Vancouver I s l a n d the w r i t e r suggests that most  of these f i l a r i a s e s simuliids.  birds  than on Vancouver I s l a n d , whereas  i n f e c t i o n s i n non-gallinaceous prevalent  numbers  dry i n t e r i o r of B r i t i s h  o r n i t h o p h i l i c s i m u l i i d s are more abundant and  C u l i c o i d e s spp.  non-gallina-  by C u l i c o i d e s only i n low  In the hot,  than i n the  immediate  f l e x i v a q i n a l i s . i n c o n t r a s t , i s not  from the C o r v i d a e ) .  Form 4 i s Mf. host  to the  Since many  specificity.  Skriabinocta specific  is' a t t r i b u t a b l e to  s p e c i f i c i t y on the part of i t s v e c t o r s .  of the passerine vicinity  b i r d s i n B. C.  are t r a n s m i t t e d  by C u l i c o i d e s . not  by  194 SUMMARY AND CONCLUSIONS •  Morphology  and Taxonomy At  l e a s t 4 ( p o s s i b l y as many as 6) species of  s p l e n d i d o f i l a r i i n e worms (Nemato.da: Onchocercidae) parasitize  the t e t r a o n i d b i r d s of B r i t i s h Columbia  and Alaska.  These a r e : l)  S k r i abinocta f l e x i v a q i n a l i s  This f i l a r i o i d  (Jones, 1961) n. comb.  occupies t h i n - w a l l e d cysts throughout the  body c a v i t y of i t s host.  This s p e c i e s ,  originally  d e s c r i b e d from a crow i n Ohio, i s more c l o s e l y r e l a t e d to S k r i abinocta petrowi Chertkova, 1946 that t o the genera Spiendidofilaria  or C h a n d l e r e l l a . where i t has been  P o s s i b l y , the b l u n t - t a i l e d , filaria  1950  sheathed M i c r o -  f l e x i v a q i n a l i s i s the same as E l l i o t t i ' s  microfilaria  placed.  (1903)  from an Ontario crow and Mf. c l a r k e i  Brinkmann,  from O n t a r i o r u f f e d grouse. The r e v i v e d genus S k r i abinocta Chertkova, 1946  comprises the f o l l o w i n g s p e c i e s : v a q i n a l i s . Sk. chitwoodae Sk. s t r i a t o s p i c u l a a l l y Sk. l i e n a l i s  Sk. p e t r o w i . Sk. f l e x i -  (Anderson, 1961) n. comb.,  ( H i b l e r , 1964) n. comb., and p r o v i s i o n (Orloff,  1947) n. comb.  With the removal of these species from i t , the genus Chandlere11a i s seen to be an a r t i f i c i a l , m u l t i g e n e r i c assemblage  of 16 s p e c i e s .  Two groups of species may deserve  separate g e n e r i c s t a t u s ; 2 other groups can be formed on a more a r t i f i c i a l b a s i s .  A more complete r e v i s i o n  i s not  195 justified  because c e r t a i n important features  species,  C. bosei  cephalic  p a p i l l a e and u t e r i n e  2)  of the type  (Chandler, 1924), v i z . . c o n f i g u r a t i o n of arrangement, are unknown.  S p l e n d i d o f i l a r i a sp. A.  A s i n g l e male was found  i n the connective t i s s u e near the oesophagus of a blue grouse. cerca  I t r e p r e s e n t s a new s p e c i e s s i m i l a r to-S.  (Lubimov, 1946) and S. caperata H i b l e r , 1964.  Possibly, (Mf.  papillo-  i t i s the adult  sp. B) of c o a s t a l 3)  of the very common m i c r o f i l a r i a  grouse.  S p l e n d i d o f i l a r i a p e c t o r a l i s n. sp.  resemble S. p a p i l l o c e r c a  and i n h a b i t  These worms  the subcutaneous  t i s s u e over the b r e a s t muscles. 4)  Splendidofilaria papillocerca  Previously  (Lubimov, 1946).  r e p o r t e d only from the U. S. S. R., t h i s species  i s now known to p a r a s i t i z e ptarmigan i n Alaska. Comparison of S. p e c t o r a l i s and S. p a p i l l o c e r c a with s i m i l a r s p e c i e s provides c o r r o b o r a t i o n (1961) c o n t e n t i o n that  the genus S p l e n d i d o f i l a r i a as he  diagnosed i t i s a homogeneous group.  Certain  p a r t i c u l a r l y of the oesophagus, r e q u i r e d i a g n o s i s be a m p l i f i e d  s l i g h t l y and that  Travassos, 1925 and S. r o t u n d i c e p h a l a relegated  f o r Anderson's  that  features, Anderson's  S. qedoe1sti  (Oschmarin, 1950) be  t o a p o s i t i o n outside the major group of 18  species. 5)  Microfilaria  lagopodis  (Haaland, 1928).  This  s p e c i e s occurs i n the blood of ptarmigan i n northern B r i t i s h Columbia  and l i k e l y  i s the embryo of S. p a p i l l o c e r c a .  196 6)  Microfilaria  sp. B.  m o r p h o l o g i c a l l y from Mf.  Not  clearly  separable  p e c t o r a l i s n. sp., t h i s  n e v e r t h e l e s s seems to represent a d i s t i n c t , new  species  (perhaps  which have escaped filaria  Splendidofilaria  detection.  and' probably  sp. A) the a d u l t s of  P o s s i b l e t h i s i s Micro-  f a l l i s i Brinkmann, 1950  O n t a r i o grouse and  form  which i s present i n  f o r which, a l s o , the a d u l t s are. unknown.  Biology On Vancouver I s l a n d f i l a r i a l p r e v a l e n t i n blue grouse adult females). encountered  i n f e c t i o n s are very  (100% of adult males; 90% of  Elsewhere,  and  i n other hosts-, they  are  less frequently. In the grouse of Vancouver I s l a n d only Micro-  filaria  sp. B and  with these Mf.  Sk.  f l e x i v a g i n a l i s occur and  s p e c i e s are hyperenzootic  sp. B has  a prevalence  adult female,  and 62%  v a o i n a l i s has  a prevalence  groups.  i n blue  grouse.  of 89% i n adult male,'72% i n  i n y e a r l i n g blue grouse; of 82%,  69%,  Sk.  sp. B  f l e x i v a g i n a l i s i s sporadic.  In the i n t e r i o r of the province Mf.  sp. B  occurs i n most species of t e t r a o n i d s but  cannot be proved  flexi-  and 38% i n these  In r u f f e d grouse of Vancouver I s l a n d Mf_.  i s e n z o o t i c , Sk.  probably  infections  at present.  In t h i s r e g i o n Sk.  this flexi-  v a g i n a l i s i s sporadic i n blue, r u f f e d , and F r a n k l i n ' s grouse,  and was  not found  i n s h a r p - t a i l e d grouse.  In blue  grouse of the s o u t h - c e n t r a l i n t e r i o r , S. p e c t o r a l i s i s hyperenzootic  i n some l o c a l i t i e s ,  enzootic i n others,  and  197 sporadic  in s t i l l  enzootic  i n spruce grouse  hyperenzootic sporadic  others.  This  filarioid  (B. C.  and  Alaska),  i n s h a r p - t a i l e d grouse  (B. C.  to  enzootic  and  to  Alaska),  and  i n r u f f e d grouse. On Vancouver I s l a n d Sk.  transmitted  flexivaqinalis is  from l a t e June to mid-August at l e a s t , by  C u l i c o i d e s u n i c o l o r group. transmitted and  i s sporadic  Mf.  sp. B on Vancouver I s l a n d i s  from e a r l y June to l a t e J u l y by Cnephia minus.  from e a r l y June to e a r l y J u l y and  i n e a r l y August, by  Simulium aureum.  The  l a t t e r black  intermediate  but  i s o f t e n l e s s abundant than  host  f l y i s the more e f f i c i e n t the  former. The Mf.  l a r v a l development of Sk.  sp. B i n t h e i r i n v e r t e b r a t e  described  f o r other  increased  temperature.  flexivaqinalis  hosts p a r a l l e l s  f i l a r i o i d s ' and  that  is accelerated  by  Apparently, development of Mf.  sp.  proceeds normally only when temperatures exceed 62°F. inferred  from young n a t u r a l i n f e c t i o n s i n chicks  prepatent p e r i o d f o r Sk.  f lexivaqina l i s  and  and  B  As  the  Mf.' sp.  B is  about 2 months, that f o r S. p e c t o r a l i s - about 6 weeks. I n f e c t i o n s with Mf. protracted  sp.  B and  Sk.  or are r e a d i l y r e - a c q u i r e d ;  S. p e c t o r a l i s are r e l a t i v e l y  The  s i m u l i i d vectors  of i n t e n s i v e f e e d i n g  adults  s h o r t - l i v e d , and  some i n d i c a t i o n t h a t immunity i s  i n the  f l e x i v a q i n a l i s ' are of there  is  conferred. of Mf.  sp. B have two  a c t i v i t y d a i l y - i n e a r l y morning  evening; these black  flies  periods and  are much more numerous  i n t r e e s than at ground l e v e l ; t h e i r a c t i v i t y  i s not  198 g r e a t l y depressed by moderate winds or at the normally  p r e v a i l i n g during  C u l i c o i d e s have only one evening on;  temperatures  t h e i r p e r i o d of abundance.  feeding p e r i o d d a i l y - from  they are more numerous i n t r e e s , but  are  late often  moderately abundant at ground l e v e l ; moderate winds suppress Culicoides  activity. The  i n the blue  hyperenzootic  nature of f i l a r i a l i n f e c t i o n s  grouse of Vancouver I s l a n d i s probably  r e l a t e d to the high p o p u l a t i o n climatic  d e n s i t y of the host  c o n d i t i o n s which favour  the breeding  h a b i t s of the v e c t o r s .  Conversely,  of B r i t i s h Columbia the  low p o p u l a t i o n  t e t r a o n i d s and  the  l e s s favourable  are r e f l e c t e d i n the  sporadic  i n the  of d i f f e r e n t  species  of c o l o u r , s i z e , and different terans  of Sk.  feeding  conditions  flexivaginalis.  the d i f f e r e n c e s i n  of d i f f e r e n t s t a t u s  seem to be the r e s u l t  a v a i l a b i l i t y to the v e c t o r s .  to  central interior  environmental  incidence  among t e t r a o n i d hosts  and  and  d e n s i t y of- the  For each species of f i l a r i o i d prevalence  directly  of  and  differential  However, the p o s s i b l e e f f e c t s  p h y s i o l o g i c a l s t a t e i n creating,,  " a t t r a c t i o n p o t e n t i a l s " for o r n i t h o p h i l i c dip-  should  be i n v e s t i g a t e d .  199 LITERATURE CITED TO END  OF PART  ONE  1.  Adams, J . R. and B e n d e l l , J . F. 1953. A high i n c i d e n c e of blood p a r a s i t e s i n a p o p u l a t i o n of sooty grouse. J. P a r a s i t . 39, Suppl: p. 11.  2.  A l i , S. M. 1956. Studies on the nematode' p a r a s i t e s of f i s h e s and b i r d s found i n Hyderabad S t a t e . Ind. J . H e l m i n t h o l . 8: 1-83.  3.  Anderson, R. C. 1954. O r n i t h o f i l a r i a f a l l i s e n s i s n. sp (Nematoda: F i l a r i o i d e a ) from the domestic duck with d e s c r i p t i o n s of m i c r o f i l a r i a e i n waterfowl. Can. J . Z o o l . 32: 125-137.  4.  Anderson, R. C. 1955. O r n i t h o f i l a r i a alaonquinensis n. sp. from Hirundo e r v t h r o q a s t e r with a r e v i s i o n of the genera P a r a m i c i p s e l l a Chow, 1939 emend. Chabaud and Choquet, 1953 and O r n i t h o f i l a r i a Gonnert, 1937. Can. J . Z o o l . 33: 107-112.  5.  Anderson, R. C. 1957. Taxonomic s t u d i e s on the genera A p r o c t e l l a Cram, 1931 and Carinema P e r e i r a and Vaz, 1933 with a proposal f o r a new genus Pseudaprocte11a n, gen. Can. J . Z o o l . 35:25-33.  6.  Anderson, R. C. 1957. The l i f e c y c l e s of d i p e t a l o n e m a t i d nematodes ( F i l a r i o i d e a , ' Dipetalonematidae): the problem of t h e i r evolution. J . Helminthol. 31: 203-224. .  7.  Anderson, R. C. 1958. On the c l a s s i f i c a t i o n of the F i l a r i o i d e a w i t h . s p e c i a l r e f e r e n c e to the F i l a r i i d a e and the S t e p h a n o f i l a r i i d a e . B u l l . Soc. Z o o l . France, 83: 144-157.  8.  Anderson, R. C. 1959. The egg and f i r s t stage l a r v a of Desmldocercella numidica (Seurat, 1920) with remarks on the a f f i n i t i e s of the Desmidocercidae. Can. J . Z o o l . 37: 407-413.  9.  Anderson, R. C. 1959. P r e l i m i n a r y r e v i s i o n of the genus D i p l o t r i a e n a Henry and Ozoux, 1909 (Diplotriaenidae: Diplotriaeninae). P a r a s s i t o l o g i a , 1: 195-307.  10.  Anderson, R. C. 1961. S p i e n d i d o f i l a r i a wehri n. sp. with a r e v i s i o n of S p i e n d i d o f i l a r i a and r e l a t e d genera. Can. J. Z o o l . 39: 201-207.  200 11.  Anderson, R. C. 1961. Study of two f i l a r i o i d nematodes, C h a n d l e r e l l a chitwoodae n. sp. from Padda o r y z i v o r a (L.) and P f ' o t o f i l a r i a . f u r c a t a Chandler, 1929. Can. J. Z o o l . 3.9: '3.17-323.  12.  Anderson, R. C. 1962. On the development, "morphology, and experimental t r a n s m i s s i o n of D i p l o t r i a e n a bargusinica ( F i l a r i o i d e a : Diplotriaenidae). Can. J. Z o o l . 40: 1175-1186.  13.  Anderson, R. C. and Chabaud, A. G. 1958. 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The l i f e c y c l e and seasonal t r a n s m i s s i o n of O r n i t h o f i l a r i a f a l l i s e n s i s Anderson, a p a r a s i t e of domestic and w i l d ducks. Can. J . Z o o l . 34: 485-525.  5.  Anderson, R. C. 1957. Taxonomic s t u d i e s on the genera A p r o c t e l l a Cram, 1931 and Carinema P e r e i r a and Vaz, 1933 with a p r o p o s a l f o r a new genus P s e u d a p r o c t e l l a n. gen. Can. J. Z o o l . 35: 93-109.  6.  Anderson, R. C. 1961. On the i d e n t i t y of A p r o c t e l l a i n b i r d s i n North America. Proc. Helm. Soc. Wash. 28: 81-82.  7.  Augustine, D. L. 1937. Observations on l i v i n g "sheathed" m i c r o f i l a r i a e i n the c a p i l l a r y circulation. Trans. Roy. Soc. Trop. Med. Hyg. 3.1" 55-60.  8.  Babero, B. B. 1953. Studies on the helminth fauna of Alaska, XVI. A survey of the helminth p a r a s i t e s of ptarmigan (Laqopus spp.) J. Parasit. 39': 538-545.  9.  Baranov, N. 1934. (Simulium reptans columbaczense i n the year 193.4). ( i n S e r b i a n : Russian summary) . V e t e r i n a r i a A r c h i v Zagreb 4: 48. • C i t e d by F a l l i s 1964 (23).  10.  B e n d e l l , J . F. 1955, Disease as a c o n t r o l of a p o p u l a t i o n of blue grouse, Dendraoapus obscurus f u l i q i n o s u s (Ridgway). Can. J . Z o o l . 33: 195-223.  11.  Bennett, G. F. 1960. On some o r n i t h o p h i l i . c bloodsucking D i p t e r a i n Algonquin Park, O n t a r i o , Canada. Can. J . Z o o l . 38: 377-389.  207 12.  Bennett, G. F. 1961. On three s p e c i e s of Hippoboscidae (Diptera) on b i r d s i n O n t a r i o . Can. J . Z o o l . 39: 379-406.  13.  Bennett, G. F. 1963. Use of P i n the study of a p o p u l a t i o n of Simulium r u q q l e s i ( D i p t e r a : S i m u l i i d a e ) i n Algonquin Park, Ontario'. Can. J . Z o o l . 41: 831-840.  14.  Bequaert, J . C. 1955-57. The Hippoboscidae or l o u s e - f l i e s (Diptera) of mammals and b i r d s . Part I I - Taxonomy, e v o l u t i o n and r e v i s i o n of American genera and s p e c i e s . Brooklyn Ent. Soc. En-tomologica Americana (n.s.) 34T36: 1-611.  15.  Boughton, D . C , Byrd, E. E., and Lund, H. 0. 1938. M i c r o f i l a r i a l p e r i o d i c i t y i n the crow. J . P a r a s i t . 24: 161-165.  16.  Chabaud, A. G. 1954. Sur le c y c l e e v o l u t i f des s p i r u r i d e s et des nematodes ayant une b i o l o g i e comparable. Ann. P a r a s i t . 29: 42-88.-  17.  Choate, T. S. 1963. H a b i t a t and p o p u l a t i o n dynamics of w h i t e - t a i l e d ptarmigan i n Montana. J . W i l d l . Mgt. 27: 684-699.  18.  Cram, E. B. 1939. R e d e s c r i p t i o n and emendation of the genus A p r o c t e l l a ( F i l a r i i d a e ) nematodes from gallinaceous birds. Proc. Helm. Soc. Wash. 6: 94-95.  19.  Dalmat, H. T. Simuliidae) v e c t o r s of C o l l . 125:  20.  Davies, D. M. and on the mating, o v i p o s i t i o n of Diptera) Can.  21.  Dutton, J . E. 1905. The i n t e r m e d i a r y host of F i l a r i a c y p s e l i Annett, Dutton, and E l l i o t t ; the f i l a r i a of the A f r i c a n s w i f t , Cypselus affinis. Thompson Yates and Johnston Lab. Rept. 6: 137-147.  22^  Edeson, J . F. B. and Buckley, J . J . C. 1959. Studies on f i l a r i a s i s i n Malaya: On the m i g r a t i o n and r a t e of growth of Wuchereria malayi i n experimentally infected cats. Ann. Trop. Med. P a r a s i t . 53: 113-119.  32  1955. The b l a c k f l i e s ( D i p t e r a : of Guatemala and t h e i r r o l e as onchocerciasis. Smithsonian Misc. 1-425. Peterson, B. V. 1956. Observations feeding, o v a r i a n -development, and adult black f l i e s ( S i m u l i i d a e , J . Z o o l . 34: 615-651.  208 23.  F a l l i s , A. M. 1964. Feeding and r e l a t e d behaviour of female S i m u l i i d a e (Diptera) . Exp. P a r a s i t 15: 439-470.  24.  F a l l i s , A. M.'an.d Smith, S. M. 1964. Ether e x t r a c t s from b i r d s and CO2 as a t t r a c t a n t s f o r some ornithophilic simuliids. Can. J . Z o o l . 42: 723-730.  25.  F a l l i s , A. M» , Davies, D. M., and V i c k e r s , M. A. 1951. L i f e h i s t o r y of Leucocytozoon simondi Mathis and Leger i n n a t u r a l and experimental i n f e c t i o n s and blood changes i n the avian host. ' Can. J . Z o o l . 29: 305-328.  26.  Gonnert, R. 1937. Zur Frage der A r t z u g e h 8 r i g k e i t von F i l a r i a mavis L e i p e r 1909. Festschrift. Bernhard Nocht zum 80. Geburtstag,- Hamburg, pp. 159-162.  27.  Guiguet, C. J . 1961. The b i r d s of B r i t i s h Columbia. (4) Upland game b i r d s , B. C. P r o v i n c i a l Museum, • Handbook No. 10.  28.  Hawking, F, 1960. P e r i o d i c i t y of m i c r o f i l a r i a e . Ind. J . M a l a r i o l . 14: 567-573.  29.  Hawking, F. 1962. M i c r o f i l a r i a i n f e s t a t i o n as an instance of p e r i o d i c phenomena seen i n host-parasite relationships. Ann. N, Y. Acad. S c i . 98: 940-953.  30.  Hawking, F. and Thurston, J . P. 1951. The p e r i o d i c i t y of m i c r o f i l a r i a e . I I . The e x p l a n a t i o n of i t s production. Trans. Roy, Soc. Trop. Med. Hyg. 45: 329-340.  31.  Hawking, F. and Webber, W. A. F. 1955. D i r o f i l a r i a • aethiops Webber, 1955, a f i l a r i a l p a r a s i t e of monkeys. I I . Maintenance i n t h e ' l a b o r a t o r y . P a r a s i t o l o g y , 45: 378-387.  32.  H i b l e r , C. P. 1961, the f i l a r i o i d s of hudsonia (Sabine, hosts i n northern  33.  H i b l e r , C. P. 1963. Onchocercidae (Nematoda: F i l a r i o d e a ) of the American magpie, Pica p i c a hudsonia (Sabine) i n n o r t h e r n Colorado. Ph.D. T h e s i s , Colorado State U n i v e r s i t y , 189 pp.  34.  Hu, S. M. K. 1939. Observations on the development of f i l a r i a l larvae during the w i n t e r season i n Shanghai r e g i o n . Am. J . Hyg. 29: 67-74.  P r e l i m i n a r y note on some of the American magpie Pica p i c a 1823) and t h e i r i n t e r m e d i a t e Colorado. Wildl'. D i s . 20.  Jensen, D. E. N. 1962. The p a t h o l o g i c a l e f f e c t s of i n f e c t i o n s of Dispharvnx nasuta (Nematoda: S p i r u r o i d e a ) on the blue grouse Dendragapus obscurus (Say). Ph.D. T h e s i s , U n i v e r s i t y of B r i t i s h Columbia, 113 pp. Menon, T. B. , Ramamurti, B. , and Rao,-'D. S. 1944. Lizard f i l a r i a s i s . Tr. Roy. So.c. Trop. Med. Hyg. 37': 373-386. Munro, J . A. and Cowan, I. McT. 1947. A review of the b i r d fauna of B r i t i s h Columbia. B. C. P r o v i n c i a l Museum, S p e c i a l P u b l i c a t i o n No. 2, 285 pp. Nelson, G. S. 1961. On Dipetalonema manson-bahri n. sp., from the s p r i n g - h a r e . Pe'detes. s u r d a s t e r l a r v a l i s . with a note on i t s development i n •• fleas. J . Helminthol..35:. 143-160. Nelson, G. S. 1962. Dipetalonema reconditum ; ( G r a s s i , ,1889) from the dog. w i t h a ; note on i t s " development i n the f l e a Ct-enocephalides f e l i s and the louse Heterodoxus s p i n i g e r . J. H e l m i n t h o l . 36: 297-308. Nelson, G„ S. 1963. Dipetalonema d r a c u n c u l o i d e s (Cobbold, 1870), from the dog i n Kenya: with a note on i t s development i n the l o u s e - f l y , Hippobosca l o n o i p e n n i s . J. Helminthol. 37: 235-240. N i c h o l a s , W, L. 1952. The bionomics of C u l i c o i d e s a u s t e n i . v e c t o r of Acanthocheilonema perstans i n the r a i n - f o r e s t of the B r i t i s h Cameroons, t o g e t h e r with notes on C. grahamii and other s p e c i e s which may be v e c t o r s i n the same area. Ann. Trop. Med. P a r a s i t o l . 47: 187-206. Odetoyinbo, J . A. 1960. Biology o f ' S p i e n d i d o f i l a r i a q u i s c a l i (von Linstow, Onchocercidae). Ph.D. Thesis,-Iowa State Univ., 142 pp. Odetoyinbo, J . A. and Ulmer, M. M i c r o f i l a r i a 1 p e r i o d i c i t y of q u i s c a l i (von Linstow, 1904) Onchocercidae). J. Parasit.  J . 1960. Spiendidofilaria n. comb.(Nematoda: 4.6, Suppl: pp. 18-19-.  Raghavan, N. G. S. and David, A. •.1955. ' F i l a r i a l i n f e c t i o n i n the grey p a r t r i d g e , F r a n c o l i n i i s pondicerianus. B u l l . N a t l . Soc. India M a l a r i a (Delhi) 3: 96.  210 45.  Raghavan, N. G. S. and M i s r a , B. G. . p r e l i m i n a r y note on experimental of avian m a l a r i a and s a u r o p s i d a l i n C. f a t i g a n s Weid. 1828. Ind. 3: 243-247.  46.  Rempel, J . G. and Arnason, A. P. 1947. An account of three successive outbreaks of the black f l y Simulium arcticum. a s e r i o u s l i v e s t o c k pest i n Saskatchewan, S c i e n t . A g r i c . 27: 428-445.  47.  1949. A infections filari.asis J. Malariol.  .Robinson, E. J , , J r . 1955. A d e s c r i p t i o n of attempts to i n f e c t mosquitoes with avian f i l a r i a l worms. J . P a r a s i t . 41: 176-178.  48.  Robinson, E. J . , J r . 1955. Observations on the e p i z o o t i o l o g y of f i l a r i a l i n f e c t i o n s i n two s p e c i e s of the avian f a m i l y Corvidae. J . P a r a s i t , 41: 209-214.  49.  Schacher, J , F. 1962. Morphology of the m i c r o f i l a r i a of Br.uqia pahanoi and of the l a r v a l stages i n the mosquito. J , P a r a s i t , 48: 679-692.  50.  Schacher, J . F. 1962. Developmental stages of Brugia pahanqi i n the f i n a l host. J . P a r a s i t . 48: 693-706.  5.1.  S c o t t , J . A. and Cross, J . B. 1946. The r a t e of growth and m a t u r i t y of Litomosoides c a r i n i i . a f i l a r i i d of the cotton r a t . J . P a r a s i t , 31, Suppl: p. 16.  52.  Webber, W. A. F. and Hawking, F. 1955. Experimental maintenance of D i r o f i l a r i a repens and D..immitis i n dogs. Exp. P a r a s i t . 4: 143-164.  53.  Weeden, R. B. 1959, The ecology and d i s t r i b u t i o n of ptarmigan i n western North America. Ph.D. T h e s i s , U n i v e r s i t y of B r i t i s h Columbia, 247 pp.  54.  Wolfe, L , S. and Peterson, D. G. 1960. Diurnal behaviour and b i t i n g h a b i t s of black f l i e s ( D i p t e r a : S i m u l i i d a e ) i n the f o r e s t s of Quebec. Can. J . Z o o l . 38: 489-497.  55.  Woo, P. T. K. 1964. A study on the blood Protozoa of blue grouse on Vancouver I s l a n d . M.Sc. T h e s i s , U n i v e r s i t y of B r i t i s h Columbia, 71 pp.  56,  YBarra, G. A. 1948. E s t u d i o monografico de nemato.dos p a r a s i t o s de l o s aves de Mexico. T h e s i s , Mexico. C i t e d by Anderson 1957 ( 5 ) .  211 APPENDIX Appendix  Status  Table I.  Sex  Occurrence of f i l a r i a l i n f e c t i o n s i n blue grouse and r u f f e d grouse from c o a s t a l B r i t i s h Columbia.  l) Dendraoapus obscurus Adult II II  ti II II II II II  n  II  II  n n  II II II II II II  II  ti II  n  ti II II II II  II II II  n  ti ii II  n II  ti  Sk. f l e x . Date ' Mf. Ad.  Locality  Mf. sp. B  fuliqinosus  m Quinsam, V . I . m ti m m tt m ti f #24 m ti f #25 m f ti m n f ti m ti m m m m m m m ti f #4 n m f , it m ti f .M m ti m m m m Nanaimo Lks., V . I. it m ti f ti m it f it f it. f m m m m ti f #3 II  II  II  II M  II  II  II  II II  II  II  H  II  II II  II  II  —0 2/6/58 0 6/6/58 , + 21/5/59 + • + 7/59 + 7/8/59 7/8/59 o. + ' + 8/59 + 7/8/50 0 + ... + 8/59 + 8/59 -+ 8/59 , + 0 7/8/59 + + 13/9/59 + ' • + 13/9/59 13/9/59 + + 13/9/59 3/60 0 5/5/60 0 + 5/5/60 + 5/5/60 0 0 0 27/7/61 - 18/7/63 + 18/7/63 18/7/63 18/7/63 ' + -0 19/7/63 .+ 19/7/63 ++ 19/7/63 + 22/8/63 + .+ 18/3/58 :  5/4/60 5/4/60 0 6/6/60 -o 9/7/60 o 28/8/60 + 31/8/60 12/5/61 + 1.2/5/61 + 20/5/61 + 24/5/61 3/7/61  -  f + +  •0 + + + + + + '  0  + .+ -r + + + + + + +  0  ++ +  +  .+ •  +  0 0 0 0  0  .+ .+ .+ + +  + + + + +  0 +  . 0  (Appendix Table I continued - 1)  Status  Sex  Adult  m m m  II  II  n  f f  II II II II II II II  ti II II II II  n II  n n II  n n II II . II  II II  n  II II II II  n  ti  II II II II  ti II  »*II II  YrIng. II II II  n  m m m m m f  m m m f f  m m m m f f f f  m f  m m m m m f  m m m m £  m f f f f f  m m m m f  m  Locality  Date  Nanaimo Lks.  6/6/62  II  #7 #8  n  19/6/62 19/6/62  19/6/62  21/6/62 II 8/6/64 . II 10/6/64 II 13/6/64 II 15/6/64 II 19/6/64 II 19/6/64 II 21/6/64 II 22/6/64 II 26/6/64 II 2/7/64 II 2/7/64 II 14/6/64 n 14/6/64 n 16/6/65 II 16/6/65 n 18/6/65 II • 3/7/65 II 3/7/65 n 5/7/65 6/7/65 " II 7/7/65 n 7/7/65 N.W. Bay, V . I . 3/6/65 n 3/6/65 n 3/6/65 n 3/6/65 n • 3/6/65 Courtenay, V.I. 21/3/64 II 9/4/64 II •11/4/64 ti 11/4/64 ii 12/4/64 II 12/4/64 n 14/4/64 II 14/4/64 n 15/4/64 II 6/5/64 n 28/5/64 ii 12/6/64 Quinsam 5/6/58 ii 20/5/59 II 3/60 II 3/60 n 31/8/63  Mf.  Ad-  B  + + +  +  +  rr + —  +++ ++ +  •++  ++ ++ + +.++ +++ + .+++ +++ +  t  + +  -—  +  —  0  —  + +  —  0  —  — —  ++ .++ — — —  +++  — —  ++ .+  -_  0  —  0 0 0  — — — —  —  +  0  + +  0 +  0 + + + + + + +  — — — —  0'  —  0  +  0  — —  + + +  ++ + +  +  —  0  .++ .+.+ .+ +++ + + +  + + ++ +  +  — — — —  0 —  + +  0  ++ + +  0  +  _  +++  0 0  —  0  .+ +  0  0 0 0 -  + +  •1-  +  213 (Appendix  Status  Sex  Locality  Yrlng.  m  Nanaimo Lks. ti  m f f f  "  m m  II II  #33 Courtenay  f  H  f f  Juvenile ca,50 juvs •**  Date  II  f  n  -f  Quinsam Nanaimo Lks. II  0  ca.40 juvs  Quinsam  2) Dendraqapus obscurus Adult  f  II  n  II  m  Juvenile n  II  f f  II  n  3) Bonasa umbellus Adult  m m f f f f f  m f  Legend:  Sk. Mf.  f lex. Ad-  Mf.  B  0  0  0 0  0 0  0  0  0 0 0  -  0  0  0  •0 0  0  0  0  22/8/60  +  +  0  4/9/60 22/8/60 4/9/60 4/9/60  +  0 0 0  0 0 0 0  • 0 0 0  0 0  0 ' 0  13/9/59 13/9/59 5/4/60 1/9/60 30/11/60 11/6/62 20/6/64 21/6/64 17/4/64 10/5/64 10/5/64 21/8/59 30/11/60 1/6-31/8 1958-61 10/6-21/7 1958-61  .++  +  + ++  + +  0 0  +  +  0  +  +  -  0  + +  -  0  -+ +  +  sitkensis  Qu. C h a r l o t t e Islands  f  [  Table- I concluded)  +  brunnescens  Nanaimo Lks. Southern V. I.' Nanaimo Lks. n II II  Courtenay Sechelt Penins. Quinsam  11/6/60 20/1/61 26/6/61 1/7/65 5/7/65 6/7/65 24/6/65 27/5/62 21/7/63  +  0 0 0 0 ' 0 0  0  +•  + -  0  -- •  0 0  +  +  -  0  .  ,+  0  -  0 = not found; - = not sought; + = specimens found. = numerous ( f o r m i c r o f i l a r i a e , about 100 per smear). +++ = over 200 m i c r o f i l a r i a e per smear. +  +  * The male S p i e n d i d o f i l a r i a  sp. A was found i n t h i s  bird.  ** Some shot, most captured and maintained up to 2 y e a r s .  Appendix Table I I . Occurrence of f i l a r i a l i n f e c t i o n s i n g a l l i f o r m b i r d s of c e n t r a l B r i t i s h Columbia.  Status l)  Sex  Locality  Dendragapus obscurus  Adult " " " " " • " " " " " " " " " " " " "  m m m m m m f f f f f m f f f #9 f #49 m #15 m m m  Date  Mf. sp. B o r * pectoralis  S. p e c t o r a l i s Ad. or Larvae  Sk. Mf.  flex. Ad.  pallidus  Trout Ck. 11  " " " " " " " Oyama " Kamloops Cherry Ck. Keremeos Penticton Kelowna Oyama Clinton " "  5/10/57 5/10/57 5/10/57 5/10/57 5/10/57 30/5/59 24/7/59 1/9/59 3/8/60 30/7/58 30/7/58 28/7/58 30/7/58 31/7/58 15/5/59 14/5/60 4/8/60 18/9/60 18/9/60 18/9/60  +++ +++ +++ + ++  0 0 0 0 0 0 0 t  +  + +  + ++ + ++  Q, 0 0 0 0 0 0 0 0 ' 0 0 0  -  — —  0 0 0 0 0 0 0 0 0 0 0 0  _  _  _ — —  0 0 _  —  _  +  +  +  0  0  0 + 0 0 0  —  0  —  _ —  (Appendix Table II continued  Status Adult n  11  II 11 11  II  Sex f f -  f m Yrlng. II f ii m 2 juv's** 5 m Juvenile ii f n f ii f 11 f ii m 2 juv's m m Juvenile II m 5 juv's Juvenile m n  ii  11  1!  f  Locality Clinton n  Ashcroft ii  II II  Gary Ranch n  Hihuna Lk. Oyama II  ti  Aspen Grove Oyama ii  II ii it  II n  Kamloops II  Cherry Ck. • Trout Ck. n  n  Date  - l)  Mf. sp. B o r * pectoralis  18/9/60 26/9/60 17/9/60 17/9/60 17/9/60 17/9/60 17/9/60 17/9/60 18/9/60 30/7/58 30/7/58 30/7/58 27/7/58 30/7-26/8/58 3/7/58 30/7/58 30/7/58 30/7/58 27/8/58 27/8/58 29/7/58 29/7/58 30/7/58 2/7-2/8/59 24/7/49 24/7/59  •t t + + 0 0 -r  + t 0 t •t  0 0 0 0 0 0 0 0 0 0 0 0 0 0  s. p e c t o r a l i s Ad „ or Larvae _  —  —  -  -  + 0 0 0 0 +  + 0 + 0 0 + +  Sk.  . fie:  M f .  + + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0  -— — —  -— — — —  _ — — —  0 0 0 0 0 0 0 _  —  _ _ _  (Appendix Table I I continued - 2)  Status Juvenile  Sex 3ex m f m f m m m m m f f m f m  f  2)  Locality  Date  Trout Ck.  11/8/59 11/8/59 19/8/59 19/8/59 1/9/59 3/8/60 12/9/59 25/9/60 17/9/60 17/9/60 17/9/60 17/9/60 17/9/60 26/9/60 12/9/59 25/9/60  II  II  ii 11  ii  Cache Ck. ii  Savona ii  Clinton ti ii ii  Lone Butte Riske Ck.  Dendraqapus obscurus  Adult ii  Juvenile II  f m _f _  m  Mf. sp. B o r * pectoralis 0 0 0 0  +  0  + + +  0 .  +  0 0 "0 0 0  S. p e c t o r a l i s Ad. or Larvae  + ++ ++ +  ?  _ — — —  _ —  0  -  Sk. Mf.  flex. Ad.  0 0 0 0 0 o • 0 0 0 0 0 0 0 0 0 0  _  0 0 0 0 0 _ _ _ _ _  _  0  richardsoni  Cranbrook E. Kootenays . . New Denver Cranbrook  9/60 12/10/58 7/11/64 9/60 9/60 9/60 9/60  0 + ++ 0 0 +  0 0  0 + 0 0 0 0 0  0 0  (Appendix Table I I continued - 3)  Status 3)  Sex  Locality  Canachites canadensis  Adult  4)  f m f m m m m m f m  Juvenile 11  m f f m f f m f m f m m m  Mf- sp. B o r * S. p e c t o r a l i s pectoralis Ad, or Larvae  Sk. Mf.-  flex. Ad.  franklini  Dog Creek Meadow Lk. Fawn Nation R. 11  Bridge Lk. 11  F t . S t . James Nazko Cold F i s h Lk.  Bonasa umbellus  Adult  Date  23/9/58 /58 23/9/60 24/9/60 24/9/60 25/9/60 25/9/60 25/9/60 28/9/60 10/61  0 0 + + t + + 0 0 (+)  19/6/59 24/9/60 24/9/60 28/9/60 23/9/60 24/9/60 25/9/60 25/9/60 26/9/60 26/9/60 30/7/58 1/9/59 12/9/59 20/9/59  0 0 0 0 0 0 0 0 0 0 0 + 0 t  0 0  - - -  +  0 0 0 - -0 0  0 0 -  —  + +  -  - 0 0  —  •  —  -  -  0 0 0 0 0 0 0 0 0 0 0 0 0 0  0 0 +  affinis  ' Trout Ck. Lac l a Hache " Quesnel Lk. Cache Ck. Ft. St. James Gary Ranch Back V a l l e y Bridge Lk. Cherry Ck. Trout Ck. L i t t l e Fort 100 Mi. House  0 + 0  (Appendix Table  Status 5)  Sex  Bonasa umbellus  Juvenile 11  6)  11  Pedioecetes  Perdix  Adult Juvenile 8)  Mf. sp. B o r * .S. p e c t o r a l i s pectoralis Ad. or Larvae  Sk. Mf.  f lex, Ad,  24/9/60 24/9/60  0 0  phasianellus  columbianus  Lac du Bois Dog Ck. " Kimberly 70 Mi. House  13/9/58 23/9/58 23/9/58 29/9/60 9/61  + t 0 0 0  17/9/61 17/9/61 17/9/61  0 0 0  14/11/61  0  + +  0 0  0 0  0 0  0 0 0 0 0  0 0 0 0  perdix f f m  Lophortyx  Adult  Date  - 4)  phaia  Cranbrook  m f  Adult m ". f Yrlng. m Juvenile m 4 (?)juv's-. 7)  Locality  I I continued  m  Cache Ck. 11  0 0 0  californicus Summerland  0  0  (Appendix Table I I concluded)  St atus  9)  Sex  Locality  Date  Mf. sp. B o r * pectoralis  S. p e c t o r a l i s Ad. or Larvae  Sk. Mf.  flex, Ad,  A l e c t o r i s qraeca  Adult  f. f m f m 6 -juv's 4m/2f  Cache Ck.  Wallachin Cache.Ck.  25/9/60 25/9/60 26/9/60 26/9/60 14/10/60 12/9/59  0 0 0 0 0 0  0 0 0 0 0 0  * M i c r o f i l a r i a sp. B cannot be s a t i s f a c t o r i l y d i f f e r e n t i a t e d from Mf. p e c t o r a l i s , T h i r d - s t a g e larvae which developed from D. p. p a l l i d u s #9, #15, and #49 s t r o n g l y i n d i c a t e d i n f e c t i o n s with Mf. sp. B. ** Only those j u v e n i l e s Note:  See Appendix  collected  a f t e r l a t e J u l y are i n c l u d e d i n t h i s  Table I f o r e x p l a n a t i o n of symbols.  table,  Appendix Table I I I .  Status l)  Sex  .Locality  Occurrence of f i l a r i a l i n f e c t i o n s i n t e t r a o n i d s of northern B r i t i s h Columbia and southern Alaska.  Date  Mf. lagopodis  S. pectoralis  S. p a p i l l ocerca  15/10/60 2/10/63 26/10/63  -  + + +  0 0 0  1/5/57 25/6/57 1/7/57 11/5/57 14/7/57 19/7/57 14/7/57 20/7/57 20/7/57 25/7/57 1/8/57 1/8/57 1/8/57 8/8/57 18/8/57 18/8/57 25/8/57 25/8/57 25/8/57 25/8/57 25/7-1/9/57  +  -  — —  —  —  -  — —  -  — —  Canachites canadensis a t r a t u s - Alaska  Juvenile n H  f m m  Mi. 176. Glenn Hwy. Taylor Hwy.* Kenai P e n i n s u l a  2) Lagopu s l a g opus (?) albus Adult F i s h Lk. m m K e l s a l l Lk. m Mi.75 Haines Road " m Mi.-72 Haines Road Mi.75 Haines Road m _ ii f f Mi.72 Haines Road ii m f Mi.75 Haines Road ii m ii f " m K e l s a l l Lk. f" Mi.75 Haines Road m Mi.76 Haines Road 11 m Mi.70 Haines Road ii m Atlin m Boulder Ck. " ii ii f II m 17 j uv s • ca. Mi.75 Haines Rd. 1  + + + +  0 0  o0 0 t 0  +  0 0 0 0 0 0 0  -  — —  — — —  — — —  _ _  _ — — —  _  -  —  -  (Appendix Table I I I concluded)  Status  Sex  Locality  3) Laqopus mutus Adult m K e l s a l l Lk. II  II  £  3 adults  f m m  Holum Ck. Mi.71 Haines Road  4) Laqopus leucurus 3 adults m K e l s a l l Lk. Adult A-tlin f 11 .£ Boulder Ck. m 11 m 3 adults.. Mi.64 Haines Road Adult Richardson Hwy. Alaska 3 juv's Mi.64 Haines Road 5) Pedioecetes p h a s i a n e l l u s caurus 7 adults - . nr. SI ana-, Tok Hwy. Adult " f Richardson Hwy. " m McGrath  *  This may be C.. c.  Note:  Date  17/5/57 17/5/57 29/5/57 15/6/57 15/7/57 18/5/57 25/8/57 25/8/57 25/8/57 25/8/57 7/9/57 10/12/63  Mf. laqopodis  S. p a p i l l ocerca  0  +  + 0 + +  0 0 0 0  0 0 0  •+  0 0 0 0 0 0 0  7/9/57 5/10/61 5/10/61 9/12/63 15/12/63  canadensis.  See Appendix Table  S. -pectoralis  I f o r explanation of symbols.  222 Appendix Table IV.  Adult non-gallinaceous b i r d s * of B r i t i s h Columbia examined for m i c r o f i l a r i a e m i a s 1959-62.  Species Accipiter striatus A c c i p i t e r 'coooeri Falco s p a r v e r i u s Rallus l i m i c o l a Porzana C a r o l i n a F u l i c a americana Charadrius v o c i f e r u s Capella qallinaoo A c t i t i s macularia Columba f a s c i a t a Zenaidura macroura Otus a s i o ' Bubo v i r o i n i a n u s C h o r d e i l e s minor Chaetura vauxi Selasphoru's r u f u s Meqacervle alcvon Colaotes c a f e r Drvocoous p i l e a t u s Sphvrapicus v a r i u s Dendrocopus v i l i o s u s Dendrocopus pubescens Tvrannus v e r t i c a l i s Empidonax hammondi Empidonax d i f f i c i l i s . Contopus s o r d i d u l u s Nuttallorriis borealis Iridioproone b i c o l o r Stelqidoptervx r u f i c o l l i s P e r i s o r e u s canadensis C v a n o c i t t a s t e l l e r i •'• Pica p i c a Corvus corax Corvus brachvrhvnchos' N u c i f r a q a columbiana Parus r u f e s c e n s P s a l t r i p a r u s minimus Sitta carolinensis Certhia f a m i l i a r i s C i n c l u s mexicanus Troqlodvtes aedon Thrvomanes bewicki Turdus m i q r a t o r i u s (Ixoreus n a e v i u s ) * *  Vane. I s l a n d Pos. 1 1 1 1  •1 3 ]_ *•*  1  1-  1  2 1  21 1  3  2  1 1 1 2 1 2 2 2 1 1 2  2 1 7 4  1  2 1 5 3  2  2 4  1 2 3 1 2 2 1  2 - 1 1  2 4  6 • 1 •- 2  Interior Neg. Pos.  1 3 1  1  1  (Appendix Table IV concluded)  Vane. I s l a n d  Interior  * Names from C h e c k - l i s t ' of North American B i r d s , Am. O r n i t h o l . Union, 5th ed., B a l t i m o r e , 1957 ( c o r r e c t e d a c c o r d i n g to Auk, 79: 493-4. 1962). ** From Vancouver. Aproctella  stoddardi  a d u l t s ; no m i c r o f i l a r i a e i n blood.  Appendix Table V.  Form 1  Form (?)1  Form 2  Form 3  Anas platyrhvnchos  Anas acuta  Fulica americana  Meqacervle alcvon  Oporornis tolmiei  5  5  5  5  10  0.5  0.5  1  6  75.6 (70.6-82.3)  78.5 (68.3-87.4)  Form 1 Host Sample Time PM (hrs.) Length (u.)  Q u a n t i t a t i v e d e s c r i p t i o n s of m i c r o f i l a r i a e from non-gallinaceous birds, i n B r i t i s h Columbia.  (?) 1 67.0 (5.7.2-79. 1)  103.3 (79.0-125.0)  MW (u.)  4.5  5  5  3  CS  1.4  2.1  5  0  {%)  267.2 (235-342) 6 3.1 (2.3-4.3)  {%)  25. 2 (24.8-26.1)  23.0 (21.4-23.8)  24.0 (23.6-24.6)  To EV {%)  34.6 (33.6-35.9)  33.8 (32.7-35.7)  33.3 (32.4-35.4)  -  35.2 (34.1-36.3)  To I B  L  {%)  55.4 (53.1-58.2)  51.6 (50.0-54.3)  50.0 • (48.1-51.1)  60.6 (57.2-64.3)  64.0 (61.2-66.1)  To I B  2  {%)  59.8 (58.3-61.7)  57.8 (55.6-59.0)  58.7 (57.7-60.6)  63.3 (60.0-67.3)  66.8 (64.4-68.2)  {%)  69.5 (67.9-71.0)  66.5 (63.5-69.3)  63.5 (62.1-64.1)  71.8 (68.6-75.0)  76.3 (74.9-78.0)  To AV [%)  82.9 (79.7-86.5)  82.8 (81.0-82.9)  82.3 (•80.1-86.2)  To NR  To R  x  38.9  24.7 (23.6-26.0)  89.8 (85.6-92.8)  (Appendix Table V continued - l )  Form 4 Dendroica auduboni  Host Sample Time PM  (hrs.)  Length ((J,) MW  Form 4  Form (?)4  Form (?)4  Pipilo .erythrophthalmus  Nuttallornis borealis  Hesperiphona vespertina  2  1  20  10  3  2  204.7 (186.0-216.0)  164.9 (150.1-169.8)  (p)  CS (ft)  0.5 (175.0-196.2)  2.9' (1.9-4.4)  4.4  24.9 (23.1-27.6)  23.7 (21.4-26.9)  To EV (ft)  36.3 (35.2-37.2)  35.8 (34.2-38.9)  To IB.,  63". 8 (61.8-67.5)  63.5 (61.5-66.2)  To I B (ft  67.0 (65.3-71.3)  66.7 (63.8-69.3)  To R (ft  77.6 (74.8-81.2)  78.7 (76.4-80.6)  75.7  To AV ft  89.3 (88.5-90.6)  89.7 (87.8-91.4)  88.6  t  190.5  6  To NR. (ft)  2  - 48 (?)  24.6  (35.7-36.2) 64.1  (66.3-69.8)  20.6  36.8 -64.1 67.6 75.0  (Appendix  Host Sample Time PM  (hrs.)  Length (|i) MW (u.)  cs  {%)  Form 5  Form 5  Nuttallornis borealis  Nuttallornis - borealis  5  "4  1  15  137.2 (132.0-146.8)  162.3 (149.5-174.7)  6  6  6.5 (4.2-9.2)  To NR  {%)  To EV  {%)  Table V continued - 2)  26.4 (24.5-27.4)  8.2 (4.8-10.0)  Form (?)5  Form 5  Form (?)5  He speriphona ve s p e r t i n a  Hvlocichla ustulata  Piranqa ludoviciana  5  10  5  48 (?)  1  4  173.8 (165.2-184.8)  125.0 (112.0-135.5)  6 6.3 (5.1-7.8)  6 6.6 • (4.7-9.3)  159.3 (141.8-177.0 6 7.6 (5.8-9.5)  25.6 (23.7-26.9)  23. 2 (21.8-24.4)  24.4 (22.5-26.0)  23.5 (22.2-25.5)  38.3 (35.8-40.1)  35.9 (33.6-37.5)  33.3 (32.7-33.9)  35.7 (32.5-37.6)  34.1 (33.2-35.5)  57.8 (55.8-61.8)  56.2 (52.6-58.2)  To IB _  {%)  60.7 (59.0-63.3)  58.2 (57.0-59.4)  55.0 (52.8-56.7)  To I E  (%)  68.8 (67.9-70.9)  65.6 (64.4-66.3)  (62.2-64.4)  65.0 (61.9-67.5)  64.0 (59.5-67.0)  76.2 (76.0-76.3)  73.8  71.4 (69.7-74.3)  70.1 (68.8-70.8)  71.0 (68.1-74.5)  • 87.4 (86.5-87.8)  84.3 (81.3-88.2)  80.8 (79.2-82.5)  84.2 (80.5-85.9)  85. 1 (82.0-87.3)  ]  To R  x  2  {%)  To AV (%)..  63.5  (Appendix Table V concluded)  Host  Form 6  Form 6  Form 6  Form 6  Form 6  Hvlocichla ustulata  Ixoreus naevlus  Sialia currucoides  Sialia mexicana  Turdus mlqratorius  Sample  5  4  2  1  1  Time PiVi (hrs.)  2  4  1  2  ?  169.7  150,7  7  -  3.3  3.5  20.5  22.6  29.1  32.4  53.5  53.3  64.4  60.3  72.7  70.4  87.8  87.8  (n)  Length  150.0 (140.0-154.5) 7  MW (u.)  146.6 (124.2-169.8) 7  (123.2-139.4) • -  4.0 (3.6-4.3)  4.8 (3.8-5.9)  (5.5-6.0)  To NR ( ft)  19.3 (18.1-20.9)  19. 5 (18.0-21.4)  (18.9-21.4)  To EV ( ft)  29.9 (27.8-32.8)  29.0 (27.5-31.6)  (27.4-29.6)  To I B  (ft)  53.7 (51.1-56.7)  55.8 (53.4-58.4)  (48.2-48.3)  (ft)  62.9 (61.2-64.2)  64.4 (62.9-66.8)  (61.7-62.3)  ( ft)  70.2 (67.8-72.5)  73.6 (72.5-75.3)  To AV ( ft)  86.9 (85.7-88.0)  89.5 (88.5-90.4)  CS (ft)  To I B To R  L  L  2  •71.8  (85.4-86.4)  228  Appendix VI. Q u a l i t a t i v e d e s c r i p t i o n s of m i c r o f i l a r i a e from non-gallinaceous b i r d s i n B r i t i s h Columbia. Form 1 Occurrence:  l ) J u v e n i l e female m a l l a r d , Anas p l a t v -  rhvnchos. 12/10/63, Ladner. acuta.  25/1/62, Pt. Roberts.  2) Adult  female p i n t a i l , Anas  (?)3) Adult male coot,  Fulica  americana. 8/2/62, Stanley Park, Vancouver. Description: Cephalic  Very short; not e v i d e n t l y sheathed.  space occupied  by 2 narrow, elongate  n u c l e i i n 3 to 4 rows, s t a i n p o o r l y to w e l l . small, often i n d i s t i n c t . hyaline.  Tail  filaria  fallisensis  attenuated,  Somatic  Excretory  Inner body very s h o r t ,  short, l i t t l e  ending i n blunt p o i n t .  nuclei.  vesicle  distinct,  u s u a l l y hooked,  S i m i l a r to m i c r o f i l a r i a  of  Splendido-  (Anderson, 1954).  Form 2 Occurrence: 6/7/59, Trout  Adult  female k i n g f i s h e r , Meoacervle  Creek.  Description:  D e l i c a t e , very narrow, l a c k i n g  Nerve r i n g , e x c r e t o r y v e s i c l e , e x c r e t o r y c e l l , very  seldom demonstrable with Giemsa.  and  T a i l tapers  sheath. anal  Form 3 Occurrence:  Adult  female M a c G i l l i v r a y ' s warbler,  t o l m i e i . 27/6/62, Nanaimo Lakes.  vesicle  slightly  to blunt t i p .  Oporornis  alcvon.  229  (Appendix VI continued - 2) Description:  Long  a n d r o b u s t ; sheath o f t e n :  extending about 50 u beyond  one e x t r e m i t y .  visible  Inner body short  but d i s t i n c t .  Caudal r e g i o n not a p p r e c i a b l y narrower than  rest  Somatic n u c l e i s t a i n w e l l .  of body.  P o s t e r i o r end of  columns of somatic n u c l e i appears truncated or c l e f t of p a i r e d a p i c a l n u c l e i 2 to 5 |i from t i p .  because  T i p of t a i l  rounded, bulbous, h y a l i n e , devoid of n u c l e i . Form 4 Occurrence:  l ) Adult female Audubon's warbler, Dendroica  auduboni. 27/6/62, Nanaimo Lakes.  2) Adult male towhee,  P i p i l o ervthrophthalmus. 27/6/62, Nanaimo Lakes.  3) Adult  male o l i v e - s i d e d f l y c a t c h e r , N u t t a l l o r n i s b o r e a l i s . 30/6/60, Nanaimo Lakes.  4) Adult female evening grosbeak,  v e s p e r t i n a . 30/3/60, Description:  Vancouver. Moderately r o b u s t , with d e l i c a t e sheath.  A n t e r i o r part of c e p h a l i c space 2.9%; Head o f t e n bent to one s i d e .  t o t a l c e p h a l i c space  Nerve r i n g r a t h e r  E x c r e t o r y v e s i c l e , i n n e r body, and anal v e s i c l e size.  Hesperiphona  Excretory c e l l  and f i r s t  from a n t e r i o r to p o s t e r i o r .  rectal  cell  9.3%.  indistinct. about same  large.  L i t t l e taper  T a i l t i p b l u n t l y rounded w i t h  t e r m i n a l n u c l e i appre.ssed to contour of t i p . Very s i m i l a r , i f not i d e n t i c a l to m i c r o f i l a r i a Skriabinocta  flexivaqinalis  of  (Jones, 1961).  Form 5 Occurrence:  l ) Adult female o l i v e - s i d e d  N. b o r e a l i s . 7/6/60, Nanaimo Lakes. 8/6/60, Nanaimo Lakes.  flycatcher,  2) Adult male N.  borealis.  3) Adult male Swainson's t h r u s h ,  (Appendix VI  230  concluded)  H v l o c i c h l a u s t u l a t a . 20/5/59, Trout Creek. western tanager, (?)5) Adult  Piranqa  (?)4) Adult  female  l u d o v i c i a n a . 27/6/62, Nanaimo Lakes.  female evening  grosbeak, H. v e s p e r t i n a . 30/3/60,  Vancouver. Description: Nuclear  Moderately short; no sheath  columns s t a i n w e l l .  Cephalic  n u c l e i u s u a l l y overlap i n n e r body. seen.  S i n g l e narrow, elongate  n u c l e a r column.  evident.  space long.  First rectal  nucleus  A  few  cell  seldom  at p o s t e r i o r end  P o s t e r i o r 6 |i of body devoid  of n u c l e i .  t i p tapers to p o i n t ; t i p o f t e n bent s h a r p l y about anal This species i s s i m i l a r to the m i c r o f i l a r i a wehri Anderson,  of  1  Tail  vesicle.  of S p i e n d i d o f i l a r i a  1961.  Form 6 •Occurrence:  Turdidae  l ) Adult  m i q r a t o r i u s . 25/6/62, Nanaimo Lakes. 27/6/62, Nanaimo Lakes. naevius.  female r o b i n , Turdus 2) Adult male r o b i n ,  3) Adult male v a r i e d thrush,  10/2/62, Vancouver.  4) Adult male Swainson's thrush,  H. u s t u l a t a . 6/7/60, Nanaimo Lakes.  5) Adult male western  b l u e b i r d , S i a l i a mexicana. 8/7/60, Nanaimo Lakes. male mountain b l u e b i r d , S. Description: nucleus-free  6)  Of medium length, with Apparently  Rectal c e l l s  1 and  fairly  P o s t e r i o r end  T a i l t i p broadly  rounded.  Creek.  long,  no sheath.  2 close together  about same s i z e ; r e c t a l c e l l s 3 and 4 c l o s e together anal v e s i c l e .  Adult  c u r r u c o i d e s . 18/5/59, Trout  c e p h a l i c space.  long i n n e r body.  Ixoreus  Moderately and near  of body almost as wide as a n t e r i o r .  231 Appendix V I I . Geographic races of the four common t e t r a o n i d b i r d s of B r i t i s h Columbia. Blue Grouse  (Dendraqapus obscurus)  D. p. s i t k e n s i s . the Alaska Panhandle  Resident on the c o a s t a l i s l a n d s  to the Queen C h a r l o t t e I s . and C a l v e r t I s .  D. p.. f u l i o i n o s u s . Alaska Panhandle  from  Resident from the mainland of the  through the c o a s t a l and western mainland  the s o u t h - c o a s t a l i s l a n d s of B. C.,  and  south to northwestern  California. D, p. r i c h a r d s o n i .  Resident from the Alaska-Yukon  southern Yukon and southwestern Mackenzie  border,  r e g i o n s , south through  n o r t h - c e n t r a l and e a s t e r n B. C. and western A l b e r t a , to Idaho, western Montana, and northwestern Wyoming. D. p. p a l l i d u s .  Resident from southern C h i l c o t i n  and  southern Cariboo r e g i o n s through the Kamloops and Okanagan r e gions south through eastern Washington to n o r t h e a s t e r n Oregon and  Idaho.  Ruffed Grouse  (Bonasa  umbellus)  B. u. yukonensis.  Resident from western Alaska across  the Yukon to southern Northwest B. u. umbelloides.  T e r r i t o r i e s and northern A l b e r t a .  Resident from southern Yukon and  northwestern B. C. south and east i n northern B. C. to c e n t r a l A l b e r t a , and east to Quebec, and south through western A l b e r t a to western Montana and northwestern Wyoming. B. u.. brunnescens.  Resident from northwestern B. C.  south i n the c o a s t a l r e g i o n to Vancouver  I s l a n d and  Vancouver.  232 (Appendix B. u. s a b i n i .  VII  Resident  concluded)from the southwestern  of B. C. south through w e s t - c e n t r a l Washington to  mainland northwestern  California. B. u. a f f i n i s . throughout  Resident  from n o r t h - c e n t r a l B. C.  most of the province east of the Coast  Washington, Oregon, and B. u. p h a i a .  south  Range, i n t o  Idaho.  Resident  from southeastern B. C.,  eastern  Washington, and northern Idaho south to e a s t e r n Oregon and s o u t h - c e n t r a l Idaho. Spruce Grouse  (Canachites  C. c.o a t r a t u s . Alaska i n v i c i n i t y  Resident i n the-;coast region, of. southern  of Kenai Peninsula and Kodiak I s .  C. c. canadensis. western  canadensis)  Resident  from extreme northern  and  A l a s k a , northern Yukon, and northern Northwest  Territories,  south through  northern B. C. to c e n t r a l A l b e r t a  and Saskatchewan, and east to c o a s t a l C. c. f r a n k l i n i .  Resident  Alaska, n o r t h - c e n t r a l B. C., out B. C.  Labrador.  from the southern Panhandle of  and western  A l b e r t a , south  through-  (except the s o u t h - c o a s t a l region) and western A l b e r t a  i n t o Washington, Idaho, western.Montana, and northwest  Wyoming.  S h a r p - t a i l e d Grouse (Pedioecetes p h a s i a n e l l u s ) P. p.. caurus.  Resident  from n o r t h - c e n t r a l Alaska  through  extreme northern B. C. to southern Yukon and northern A l b e r t a . P. p.. columbianus.  Resident  of the Coast Range, south through northern New  Mexico.  from n o r t h - c e n t r a l B. C. east south-central B. C. as f a r as  233 Appendix Biting  VIII.  i n s e c t s from the study areas.  Mallophaga Laoopoecus  spp.  Diptera 1) C u l i c i d a e Trout  Creek  Aedes canadensis  - from a e r i a l sweeps  (Theo.)  Aedes cinereus Meig.  - from a e r i a l sweeps  Aedes f i t c h i  - from a e r i a l sweeps  (F. and Y.)  Aedes stimulans (Walk.)-  - from a e r i a l sweeps  Anopheles sp.  - from a e r i a l sweeps  Nanaimo  Lakes  Aedes canadensis  - near, or b i t i n g  man  Aedes cinereus  - near, or b i t i n g  man  Aedes  - near, or b i t i n g  man  - near, or b i t i n g  man  - near, or b i t i n g  man  - near, or b i t i n g  man  - near, or b i t i n g  man  fitchi  Aedes s t i c t i c u s  (Meig.)  Aedes stimulans Aedes v a r i o a l p u s Cu.lex t e r r i t a n s  (Coq.) Walk.  C u l i s e t a incidens Mansonia  (Thorn.)  - ex grouse, not engorged  oerturbans (Walk.)  - ex grouse,  Anopheles p u n c t i p e n n i s (Say)  engorged  - from sweeps near man  2) Ceratopogonidae Trout Creek - a l l engorged Culicoides  from blue 'grouse  crepuscularis Mall,  Culicoides unicolor  group  (Coq.) group  C u l i c o i d e s sp. Nanaimo  Lakes  Culicoides  crepuscularis  group - ex grouse, abundant  Culicoides  (?)luteovenus R. and H. - ex man  engorged,  234  (Appendix V I I I continued - l ) C u l i c o i d e s obsoletus group  ex  G. p i l i f e r u s / u n i c o l o r group sp. .1 C. p i l i f e r u s / u n i c o l o r group sp. 2 Culicoides unicolor Culicoides  sp,  Culicoides  sp,  group  man  - ex  man  - ex man ex grouse, moderately ex grouse, abundant  - ex  engorged, abundant engorged,  man  Simuliidae Trout  Creek  Prosimulium dicentum D. and S.  - ex,grouse,  engorged  (2 specimens), and Prosimulium fulvum Coq.  - near  man  man  Prosimulium p l e u r a l e M a l l . Simulium  (Euslmulium)  s.. (Eu.) ( ? ) o s b o r n i s. s. s. s.  - ex grouse, engorged (2 specimens) aureum, F r i e s - ex grouse, engorged, very abundant  (Stains and -- ex grouse, engorged Knowlton) (1 specimen) - ex grouse ( l specimen)  (Eu.) sp.  (Eu.) sp. (s.) arcticum M a l l .  - ex grouse - ex man  (s.) canadense Hearle  - ex grouse, engorged (1 specimen), and man -- ex man (1 specimen)  s. (s.) venustum Sav Cnephia minus (D. and  ( l specimen)  S.)  - ex grouse,, engorged, abundant  Nanaimo Lakes Prosimulium exiqens D. and S, . Prosimulium. dicentum Prosimulium  Twinn  Prosimulium novum D. and  man  ex man - males, ex grouse ( c o l l Dr. J . F. Bendell)  pleurale  Prosimulium unicum  ex  S.  - ex  man  - ex  man  S.  (E_u.) aureum  - ex grouse, engorged, very abundant  S.  (Eu..) sp.  - near  S,  (S.) venustum  - ex grouse, engorged ( l specimen)  man  235 (Appendix V I I I concluded)S. ..(S.) S.  canadense  reared from pupa  (S.) O c c i d e n t a l e Towns.  Cnephia m i n u s  4) Tabanidae  -  ex grouse, engorged (2 specimens) ex grouse, engorged, very abundant; and near man -  Nanaimo Lakes  Tabanus sp.  near  man  Chrysops  near  man  5) Hippoboscidae  spp. -  Quinsam r e g i o n  Ornithoica vicina  (Walk.)  Qrnithomyia f r i n q i l l i n a  Neolipoptena  ferrisi  Latr.  (Beq.)  ex juv.. -blue grouse ex s e v e r a l 'species of p a s s e r i n e b i r d s ; ex yr, blue grouse; ex crow ( c o l l . Dr. J . F. . B e n d e l l , 1950-63) 6 ex j u v . blue grouse ( c o l l . Dr. J. F. B e n d e l l , 23/8/60)  ABBREVIATIONS USED IN PLATES I-VI  am  amphid  av  anal  ep  excretory  pore  ev  excretory  vesicle  9P  genital  ib  i n n e r body  nr r  l.  vesicle  primordium  - nerve r i n g first rectal  cell  ( A l l drawings made with the a i d of a m i c r o p r o j e c t o r . )  EXPLANATION OF PLATE I Adults of S k r i a b i n o c t a f l e x i v a o i n a l i s Figure  1.  Left  l a t e r a l view a n t e r i o r end of female,  Figure  2.  En face view of c e p h a l i c apex.  Figure 3.  Right  l a t e r a l view caudal  end of male.  Figure 4.  Right  l a t e r a l view caudal  end of female.  Figure 5.  Right  l a t e r a l view of r i g h t  Figure 6.  Left  Figure 7.  M i c r o f i l a r i a from t h i n  l a t e r a l view of l e f t  spicule.  spicule.  smear.  ( F i g s . 5, 6, and 7 drawn to same s c a l e . )  236  PLATE I  EXPLANATION OF PLATE I I Adult Male S p l e n d i d o f i l a r i a  sp. A  Figure 1.  V e n t r a l view of a n t e r i o r end.  Figure 2.  Left  Figure 3.  En face view of c e p h a l i c  Figure 4.  Right l a t e r a l view of caudal end.  Figure 5.  Right l a t e r a l view of r i g h t s p i c u l e  Figure 6.  Left  Figure 7.  O p t i c a l s e c t i o n of c u t i c l e .  l a t e r a l view of a n t e r i o r end. apex.  l a t e r a l view of l e f t s p i c u l e .  (Figsv 1 and 2 drawn to same s c a l e ; f i g s . 5 and 6 drawn to same scale.),  237  EXPLANATION OF PLATE I I I Adult Female S p l e n d i d o f i l a r i a p e c t o r a l i s n, Figure 1.  Right l a t e r a l view of a n t e r i o r end.  Figure 2.  En face view of c e p h a l i c apex.  Figure 3.  Right l a t e r a l view of v u l v a r r e g i o n  Figure 4.  Left  Figure 5.  V e n t r a l view of caudal end.  Figure 6.  Microfilaria  l a t e r a l view of caudal end.  from t h i n  smear.  ( F i g s . 1 and 4 drawn to same s c a l e ; f i g s . 3 and 5 drawn to same s c a l e . )  238  PLATE III  6  f  50^  EXPLANATION OF PLATE IV Adult Male S. p e c t o r a l i s Figure  1.  Left  Figure 2.  Right  Figure 3.  Left  n. sp.  l a t e r a l view of caudal end. l a t e r a l view of r i g h t l a t e r a l view of l e f t  spicule.  spicule..  (.Figs. 2 and 3 drawn t o same s c a l e . )  Fourth-stage  Larvae S. p e c t o r a l i s  n. sp.  Figure 4.  Right  lateral  view a n t e r i o r end of male.  Figure 5.  Right  l a t e r a l view of v u l v a r r e g i o n .  Figure 6.  V e n t r a l view caudal  Figure 7.  Right  end of female.  l a t e r a l view caudal  end of female.  ( F i g s . 6 and 7 drawn t o same s c a l e . )  239  PLATE IV  EXPLANATION OF PLATE V Adult S p l e n d i d o f i l a r i a p a p i l l o c e r c a Figure 1.  V e n t r a l view a n t e r i o r end of female.  Figure 2.  En face view of c e p h a l i c apex.  Figure 3.  Left  Figure 4.  Right l a t e r a l view caudal end of female.  Figure 5.  Right l a t e r a l view caudal end of male.  Figure 6.  Right l a t e r a l view of r i g h t s p i c u l e .  Figure 7.  Left  Figure 8.  V e n t r a l view caudal end of female.  Figure 9.  O p t i c a l s e c t i o n of c u t i c l e .  l a t e r a l view a n t e r i o r end of female.  l a t e r a l view of l e f t s p i c u l e .  ( F i g s . 1, 4, 5, and 8 drawn to same s c a l e ; f i g s . 6 and 7 drawn t o same s c a l e . )  240  PLATE V  EXPLANATION OF PLATE VI Microfilaria  sp. B and Mf. lagopodis  Figure 1.  Microfilaria  sp. B from t h i n smear.  Figure 2.  V e n t r a l view second-stage l a r v a of Mf.. sp. B.  Figure 3.  Right l a t e r a l view a n t e r i o r end of t h i r d - s t a g e l a r v a of Mf. sp. B.  Figure 4.  Right l a t e r a l p o s t e r i o r end of t h i r d - s t a g e l a r v a of Mf. sp. B,  Figure 5.  V e n t r a l view p o s t e r i o r end of t h i r d - s t a g e l a r v a of Mf. sp. B.  Figure 6.  Mf, lagopodis from t h i n  smear.  ( F i g s . 1, 3, 4, and 6 drawn to same s c a l e . )  241  PLATE VI  

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