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The termites of British Columbia, their structure, bionomics and intestinal fauna Jacob, John Kenneth 1938

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THE TERMITES OF B R I T I S H COLUMBIA, THEIR STRUCTURE, BIOHOMICS AND INTESTINAL by John Kenneth Jacob  A T h e s i s s u b m i t t e d f o r t h e Degree o f MASTER OF ARTS i n t h e Department o f ZOOLOGY  UNIVERSITY OF B R I T I S H COLUMBIA April  .1938  FAUNA-  I Acknowledgements In w r i t i n g these, it  t h e most p l e a s a n t  l i n e s o f my  i s w i t h a s i n c e r e s e n s e o f g r a t i t u d e t h a t I o f f e r my  the f o l l o w i n g people, p r e p a r a t i o n of the I am  whose i n t e r e s t and  thesis,  thanks to  h e l p h a v e made t h e  thesis possible.  e s p e c i a l l y indebted  to Dr.  C. M c L e a n  Head o f t h e D e p a r t m e n t o f Z o o l o g y , f o r h i s  Eraser,  a s s i s t a n c e and  en-  couragement. My w o r k was was  t h a n k s a r e due  first  s u g g e s t e d and  c a r r i e d on.  His  in  G. M.  the p r e p a r a t i o n To Mr.  wish  to e x t e n d my  G. J . S p e n c e r , by whom t h e  u n d e r whose d i r e c t i o n a l l t h e work  i n v a l u a b l e a s s i s t a n c e when t h e  seemed most d i f f i c u l t Dr.  t o Mr.  i s deeply  appreciated.  Y/atney s u p p l i e d r e f e r e n c e s o f some o f t h e  Perry  and  gave  of the F o r e s t P r o d u c t s  Laboratory,  and  duly t h a n k f u l :  i n x>reparation  preparation  graduates i n the  Mr.  p r e p a r a t i o n o f many o f t h e p h o t o g r a p h i c for i n v a r i a b l e advice  I  tests.  the f o l l o w i n g s t u d e n t s ment o f Z o o l o g y , I am  advice  drawings..  g r a t i t u d e f o r h i s a s s i s t a n c e i n the  o f t h e wood p r e f e r e n c e  tance  problem  D.  B.  Quayle f o r  p l a t e s , Mr.  i n p h o t o g r a p h y , Mr.  Depart-  Wm.  the  Cameron  J . L. McHugh f o r a s s i s -  o f t h e t a b u l a r d a t a and  Mr.  0. P.  Idle for  m a t e r i a l from Vancouver Island,. I have r e c e i v e d v a l u a b l e B. H. B.  i n f o r m a t i o n from P r o f e s s o r  S t r i c k l a n d o f t h e U n i v e r s i t y o f A l b e r t a and  L e e c h , now  at the U n i v e r s i t y of  C.  and Mr.  H.  A n d i s o n of  Hugh  California.  F o r many s p e c i m e n s I w i s h Hope, B.  f r o m Mr.  to thank Mrs.  Vicjoria.  B.  ^hacker  of  INDEX  Page  INTRODUCTION. General explanation of subject Procedure Technique  1 3 4  CLASSIFICATION. Termites,  t h e i r a n c e s t o r s and t h e i r to o t h e r i n s e c t s . Characteristics of the Isoptera Characteristics of the f a m i l i e s Key t o t h e B r i t i s h C o l u m b i a S p e c i e s  DISTRIBUTION. Geographical d i s t r i b u t i o n Factors affecting geographical PART I THE STRUCTURE OF B R I T I S H  relations  7 9 10 12 13 15  distribution  COLUMBIA TERMITES  EXTERNAL ANATOMY: A c o m p a r i s o n o f t h e t h r e e s p e c i e s The Head. Head c a p s u l e o f a l l c a s t e s a n d s p e c i e s Antennae Mouthparts Tentorium The  Thorax. C e r v i x o f a l l c a s t e s and s p e c i e s Hota Pleura Sterna Endoskeleton Legs Wings  ,  The Abdomejn. Segments, a b d o m i n a l appendages and g e n i t a l i a  17 26 29 37  6  41 42 48 52 55 1 68 72  INTERNAL ANATOMY: A c o m p a r i s o n o f t h e t h r e e s p e c i e s . The D i g e s t i v e T r a c t . General Morphology: p h a r y n x , oesophagus, crop, g i z z a r d , oesophageal v a l v e , mid i n t e s t i n e , Malpighian tubules, small intestine, large i n t e s t i n e , c o l o n , rectum and accessory glands. A l a t es gootermopsis a n g u s t i c o l l i s 77 nevadensis 81 R e t i c u l i t eriaes h e s p e r u s 81  The R e p r o d u c t i v e S y s t e m . Female. General Morphology: ovary, o v i d u c t s , v a g i n a , seminal receptacle, c o l l e t e r i a l glands, genital armature. Alatea. Z. a n g u s t i c o 1 1 i s 83 R. h e s p e r u s 85 Other castes 85 Male. General Morphology: t e s t i s , vasa d e f e r e n t i a , seminal v e s i c l e s , accessory glands, e j a c u l a t o r y duct, g e n i t a l armature. Alates. Z, n e v a d e n s i s 2, a n g u s t i c o i l i s R. h e s p e r u s Other c a s t e s The H e r v o u s Sy at em. General Morphology: 1. C e n t r a l n e r v o u s s y s t e m - b r a i n v e n t r a l nerve cord. Alates. Z. a n g u s t i c o i l i s E..hesperus. Other c a s t e s 2. S t o m o d a e a l n e r v o u s s y s t e m . Alates. 2. a n g u s t i c o i l i s R. h e s p e r u s The The The  ;  and  Ser»3e O r g a n s . H a i r s , p o r e s and o t h e r s e n s e o r g a n s . T r a c h e a l System. Spiracles, tracheal  trunks,  87 88 89 90  tracheoles.  Organs o f D i s t r i b u t i o n , C o n s e r v a t i o n and Elimination. The B l o o d "' The O r g a n s o f C i r c u l a t i o n : h e a r t and aorta The F a t Body The O e n o c y t e s The C o r p o r a A l a t a ,  92 96 97 98 99 99 101  103 103 104 104 104  PART I I . BIONOMICS L I F E HISTORY. The C o l o n i z i n g f l i g h t , s w a r m i n g a n d p a i r i n g . Fertilization. Egg L a y i n g Care o f Eggs Incubation period Biotic potential Developmental stages Embryonic* P o s t emb r y o n i c P o s t - a d u l t growth and l o n g e v i t y . Role of supplementary r e p r o d u c t i v e s C o n s t i t u t i o n of the colony D i s t r i b u t i o n w i t h i n the colony Theory o f c a s t e d i f f e r e n t i a t i o n  105 106 106 107 107 108 108 109 110 111 112 113 114  TROPISMS. ~ Heliotropism Thermotropism T h i gmo t r o p i sm Chemotropiam  118 119 119 119 120  HA33ITS * '  120  MTJS:£L ~  ENEMIES. I n q u i l i n e s and P r e d a t o r s  121  PART I I I INTESTINAL FAUNA Introduction Protozoa L i s t of protozoa o c c u r r i n g i n the three species C l a s s i f i c a t i o n of these protozoa Distribution within the digestive tract Cell Division P h y s i o l o g i c a l R e l a t i o n s h i p between P r o t o z o a and t e r m i t e Protozoan pojjulation i n v a r i o u s stages of the host F a c t o r s a f f e c t i n g protozoan content Refaunation following ecdysis E f f e c t on p r o t o z o a c a u s e d "by f e e d i n g t e r m i t e s on woods o t h e r t h a n t h o s e p r e f e r r e d by t e r m i t e s . P a r a s i t i c p r o t o z o a as a means o f c o n t r o l .  122 122 122 123 128 128 128 130 131 132 133 134  Page. PART I V . FUNGI AND THEIR RELATIQII TO TERMITES. The f u n g i ' F u n g i as a source  135 136  of food PART V.  ECONOMIC STATUS. Type o f w o r k a n d e x t e n t o f damage i n B r i t i s h Wood P r e f e r e n c e s .  C o l u m b i a 137 138  CONTROL.  139  CONCLUSION.  141  A p p e n d i x , a c o m p l e t e r e c o r d o f wood p r e f e r e n c e Tables  1-5  Bibliography  tests.  142  THE  TERMITES OF BRITISH COLUMBIA*  THEIR STRUCTURES, BIONOMICS AND  INTESTINAL FAUNA„  Introduction, Termites, the f i r s t  l i k e other s o c i a l i n s e c t s , were among  to r e c e i v e the a t t e n t i o n of entomclogists  naturalists  and  ( M a e t e r l i n c k ) , the t r o p i c a l species being pop-  u l a r i z e d more than the temperate, probably  because of  l a r g e nests* As a r e s u l t of the e a r l y w r i t i n g s on  their  termites  most people a s s o c i a t e them only w i t h the t r o p i c s . Although it  i s true that the m a j o r i t y of them are c o n f i n e d to  e q u a t o r i a l b e l t , there are some species which extend  the their  range i n t o the warmer temperate zones and a l i m i t e d number which l i v e only i n the c o o l e r p o r t i o n s of the  temperate  zones. In -She n o r t h temperate zone t h e i r northern  distrib-  u t i o n extends i n Europe s l i g h t l y n o r t h of Spain, and i n North America, south east of Lake S u p e r i o r , and on P a c i f i c coast, as f a r n o r t h as Skidegate,  the  Q,ueen C h a r l o t t e  Islands. T r o p i c a l species are f o r the most p a r t d e s t r u c t i v e s while  temperate forms although not so harmful cause c o n s i d -  erable damage both to indoor as w e l l as  to outdoor s t r u c t u r e s .  In B r i t i s h Columbia the damage caused by termites i s c h i e f l y restricted  to woodwork l y i n g c l o s e to the ground or wood apt  to become wet  and  decayed. As a r e s u l t of the i n c r e a s i n g  -2number of r e p o r t s of termite damage, i t became apparent that the bionomics of, and  i n j u r y caused by them, should  be  7  i n v e s t i g a t e d . This t h e s i s i s an attempt to supply  this  information. The problem was  a t t a c k e d by;  (a) l i s t i n g  able r e c o r d s of termites i n the p r o v i n c e ; a d d i t i o n a l r e c o r d s by personal search, and aid  of f r i e n d s , entomologists,  collectors;  a l l avail-  (b) o b t a i n i n g fefVlisting  interested citizens  the  and  ( c ) r e a r i n g through two y e a r s , samples of  e c t i o n s from s e v e r a l sources  coll-  i n l a b o r a t o r y incubators  with  c o n t r o l l e d temperature and humidity;  (d) making a compar-  a t i v e h i s t o l o g i c a l and morphological  study of a l l the  B r i t i s h Columbia s p e c i e s and of  the i n t e s t i n a l  (e) "by making a s p e c i a l  study  protozoa*  As a r e s u l t of c o n s o l i d a t i n g the c o l l e c t i o n s i n f o r m a t i o n obtained  there were found  n a t i v e to t h i s province, two a n g u s t i c o i l i s Hagen and the c o a s t a l r e g i o n and Banks and  s  found  and  to be three s p e c i e s  of them, Zooterraopsis  Z. nevadensis  Hagen, o c c u r r i n g i n  the t h i r d , R e t i c u l i t e r m e s hesperus  i n the d r i e r p a r t s of east Vancouver I s l a n d  the Okanagan V a l l e y .  o^B.. #^ -fp^ v  , --  ,  ' ^' \ tM  4  sU/v  ~3PROCEDURE AND Items (a) and  TECHNIQUE«  (b) concerning r e c o r d s of occurrence  w i l l be i n c o r p o r a t e d i n the substance so we w i l l proceed  of t h i s  dissertation,  to s e c t i o n (c) d e a l i n g w i t h the bionom-  i c s of the three s p e c i e s . The s p e c i e s Z. a n g u s t i c o l l i s was experiments  as i t i s the most common of the three s p e c i e s *  M a t e r i a l was was  used i n a l l my  obtained c h i e f l y from.decayed stumps where i t  found the l a r g e s t c o l o n i e s occur,, The  t e r m i t e s were  p l a c e d i n l a r g e g l a s s b a t t e r y j a r s w i t h p o r t i o n s of the wood and when p o s s i b l e , a p a r t of the tunnels i n which they l i v e d . The openings  j a r s were c l o s e d w i t h l i d s each having  two  i n t o which corks were i n s e r t e d . In t h i s way  i t e s themselves  term-  could r e g u l a t e the humidity w i t h i n the  ©olony. This method proved  e n t i r e l y s a t i s f a c t o r y f o r main-  t a i n i n g a l a r g e supply. For o b t a i n i n g eggs a t r e g u l a r i n t e r v a l s * and newly-emerged specimens f o r embryological s t u d i e s j other c o l o n i e s were p l a c e d i n p e t r i d i s h e s w i t h wood* moistened  from time to time to maintain the proper  humidity. A l l these c o l o n i e s were kept a t room  temperatures.  To hasten the development of seventh i n s t a r i n d i v i d u a l s about to become a l a t e s they were p l a c e d i n an i n c u b a t o r kept a t temperatures  r a n g i n g between 65' and 80*E , w i t h  the r e l a t i v e humidity maintained was  e  a t about 80 percent. I t  soon found simpler and e a s i e r to r a i s e c o l o n i e s of  many i n d i v i d u a l s *  than those of o n l y a  few.  TECHNIQUE. In making the morphological and h i s t o l o g i c a l studies  the f o l l o w i n g methods were^found the most satis°  f a c t o r y . In the p r e p a r a t i o n specimens were a n a e s t h e t i z e d  of h i s t o l o g i c a l m a t e r i a l , w i t h cyanide gas,  the i n t e r -  segmental membranes punctured w i t h minuten pins to penetration  "  allow  of the f i x , immediately immersed i n Bouin's  picro-formol,  washed i n 50 percent a l c o h o l and  dehydrated,  c l e a r e d i n x y l o l i f the o b j e c t v/as s m a l l , or i n cedar o i l i f l a r g e , and  i n f i l t r a t e d w i t h p a r a f f i n kept a t 54*0. This  procedure was  e x c e l l e n t f o r s e c t i o n s of whole i n s e c t s where  minute c e l l u l a r examination was  not d e s i r e d . Eor  cytolog-  i c a l d e t a i l s the  i n s e c t s were cut i n t o three p a r t s v i z . ,  head, thorax and  abdomen, f o l l o w i n g anaesthetization«,after  'which the procedure was  the same. Sections  were s t a i n e d  w i t h Heidenhain's or H a r r i s * haematoxylin and Mallory s r  t r i p l e s t a i n , and In a d d i t i o n to the  mentioned, b u t y l a l c o h o l was  methylene two  proportions  blue.  c l e a r i n g agents above  found to be e x c e l l e n t , as i t  does not harden c h i t i n as much as used i n v a r y i n g  eosin,  the other two.  w i t h e t h y l a l c o h o l as  ( t h i s s e r i e s f o l l o w s f i x a t i o n of the t i s s u e ) :  It  was  follows  35%  ethyl alcohol — ~ — —  900c.c. of 45%  ethyl alcohol  100c.c. of n-butyl 800c.c. Of 62%  ~ — —  — — -  °  200c.c. - - - - - - 2 h r s . 350coC — - —  —-  250c.c. of 100%  --.-—  -----------  4 hrs.  550c.o, ------- 6 h r s . to days. 750c.c. ------- 6 h r s . to overnightrwith  n-butyl  a l o n e : two Following  to a 1:2  one  change.  changes a t i n t e r v a l s of s e v e r a l hours. t h i s the  mixture of n-butyl  t i s s u e s were then t r a n s f e r r e d and  p a r a f f i n i n which they  remained f o r s e v e r a l hours f o l l o w e d changes of p a r a f f i n . With the use t r a t i o n may  - 1 hr,  a l c o h o l —------------ approx. 2 h r s .  650c.c. of 11% 450c.c. of 90%  30 min.  by  three or  four  of b u t y l a l c o h o l  be g i v e n as much as one  or two  infil-  days f o r  completion. In f i x i n g embryos the eggs were punctured w i t h minuten pins w h i l e immersed i n Bauin's. A f t e r  complete  f i x a t i o n i t was  removed  found that i f the chorion was  b e t t e r dehydration and obtained. Also  i t was  i n f i l t r a t i o n of the p a r a f f i n found u s e f u l to s t a i n the  was  embryos  w i t h a 95% a l c o h o l s o l u t i o n of b a s i c f u c h s i n so that  they  could be more e a s i l y o r i e n t a t e d i n the p a r a f f i n b l o c k s . For  examination of s c l e r i t e s the  c l e a r e d i n a 10% use  specimens were  s o l u t i o n of potassium hydroxide. With the  of a hot mixture of 30% a l c o h o l plus 5 .percent g l a c i a l  a c e t i c a c i d the specimens could be distended  to show more*  -6In s t u d y i n g l i v e p r o t o z o a a common s a l t of i 5 9 % w s  excellent.  Celled  solution  s l i d e s s e a l e d w i t h petrolatum  kept the p r o t o z o a i n a f a i r l y a c t i v e s t a t e f o r a much longer time than any  other method.  TERMITES OF THE PAST AND  THE ANCESTORS OF PRESENT DAY  SPECIES.  From i n f o r m a t i o n obtained from f o s s i l r e c o r d s ,  term-  i t e s would seem to have o r i g i n a t e d at l e a s t as f a r hack as Mesozoic, although p e r i o d (Wheeler  no r e c o r d s have "been obtained  *28).  Holmgren# has  the  from t h i s  shown that an A u s t r a l i a n  s p e c i e s , Mastotermes d a r w i n i e n s i s , must be d e r i v e d from the P r o t o b l a t t o i d e a , a group which became e x t i n c t i n the Permian. T h i s super-family  of the order P r o t p r t h o p t e r a ,  containing  many c o c k r o a c h - l i k e f o s s i l i n s e c t s taken from s t r a t a from the Carboniferous two branches, one roaches,  ranging  to the T r i a s s i c epochs, gave r i s e to  t e r m i n a t i n g i n the f o s s i l and present  the P a l a e o p t e r a  day  ( P r o t o b l a t t i d a e and B l a t t i d a e ) ,  the  other i n the I s o p t e r a , p r i m i t i v e r o a c h - l i k e i n s e c t s commonly termed t e r m i t e s . Present  day forms such as the p r i m i t i v e species  Mastotermes d a r w i n i e n s i s and Archotermopsis wroughtoni of the North-Y/estern  Himalayas, have many c h a r a c t e r s which p o i n t back  to ancestors  c l o s e l y r e l a t e d to those of the roaches of today.  The  f o l l o w i n g c h a r a c t e r s are d i s t i n c t l y b l a t t o i d : i n the former  s p e c i e s the presence of the anal lobe i n the h i n d wing, wing venation which i s l i k e that of the P r o t o b l a t t o i d s , 5 - j o i n t e d #  '09-'13. Handl.  Termitenstudien,  4 parts.  See Wheeler's b i b l i o g r a p h y .  K.  Svensk.  Vetensk.  t a r s i , spined t i b i a e , absence of f r o n t a l gland, presence of three p a i r s of d i s t i n c t  gonapophyses  (Crampton  '20) and the  p e c u l i a r s t r u c t u r e of hypodermis c h a r a c t e r i s t i c of roaches (Jucci  '24); i n the l a t t e r  s p e c i e s , the s t r u c t u r e of the  d i g e s t i v e and r e p r o d u c t i v e systems of both sexes, presence of caeca i n the midgut, the eight M a l p i g h i a n v e s s e l s (the same number as i n young roaches), the hypopharynx, the resemblance of v e s i c u l a e s e m i n a l i s of male to the mushroom gland of roaches and spined t i b i a e  (Imms '19-).  There are f i v e orders of i n s e c t s c l o s e l y r e l a t e d to the I s o p t e r a , which possess some t e r m i t e c h a r a c t e r s ; ( l ) the B l a t t i d a e i n which the s p e c i e s Cryptocercus p u n c t u l a t u s , the l a r g e , p r i m i t i v e , w i n g l e s s , brown roach, l e a d s a somewhat social l i f e  i n decaying wood and has a protozoan fauna showing  a very s t r i k i n g resemblance to the fauna of some t e r m i t e s . (2) the Embioptera i n which maternal care of the young l a r v a e and ova i s s t r o n g l y e x h i b i t e d by the females, ( 3 ) the Dermapt era of which  fforficula  a u r i c u l a r ! a shows maternal care of i t s  young, (4) the Zoraptera which bear a g r e a t e r resemblance to t e r m i t e s than the above mentioned o r d e r s , l i v e under dead bark or wood (sometimes i n the v i c i n i t y of t e r m i t a r i a ) , may may  or  not shed t h e i r wings and the apterous a d u l t s are b l i n d ,  and f i n a l l y ,  ( 5 ) the order G r y l l o b l a t t i d a e which Crampton  claims to be a l i n k between  the Dermaptera and I s o p t e r a .  (*15)  -9CLA.3SIEICATI0N. The  t e r m i t e s c o n s t i t u t e a separate  order, the  I s o p t e r a B r u l l e , s o - c a l l e d "because a l a t e forms have the f o r e and h i n d wings of n e a r l y equal  size.  T h i s small order c o n s i s t s  of 152 genera and subgenera and 1978 d e s c r i b e d species  (Synder  '35), i n c l u d i n g v a r i e t i e s and subspecies, mostly from the warmest p o r t i o n s of the t r o p i c s ; about a t h i r d a r e from A f r i c a , North America has 56 s p e c i e s , two of which are i n t r o d u c e d . There a r e f o u r species i n Canada^<»oPt^^'^\We^te'tusaBr>tvsVVC£l«»*fcvai  Kl  The by  I s o p t e r a may be d i s t i n g u i s h e d from other  orders  the f o l l o w i n g c h a r a c t e r i s t i c s#; S o c i a l and polymorphic i n s e c t s l i v i n g i n l a r g e  communities, composed of winged and / or brachypterous and / or apterous r e p r o d u c t i v e forms together with numerous sterile,  s o l d i e r s and workers.  b i t i n g type: l i g u l a 4-lobed. membranous, superposed f l a t capable  apterous,  Mouth p a r t s of the t y p i c a l  Tf/ings very  s i m i l a r , elongate and  over the back when at r e s t and  o f being shed by means of b a s a l f r a c t u r e s : a n t e r i o r  v e i n s s t r o n g l y c h i t i n i z e d , r e g u l a r cross v e i n s wanting and an i r r e g u l a r network o f t e n present always 4-jointed„  (archedictyon).  T a r s i almost  C e r c i short or very s h o r t : g e n t a l i a wanting  or rudimentary i n both sexes (except  i n Mastotermes).  morphosis s l i g h t or absent. The #  order may be d i v i d e d i n t o f i v e f a m i l i e s which Adapted from Imms, Textbook of Entomology '25.  Meta-  -10are separated as f o l l o w s : 1. Mastotermitidae  S i l v e s t r i '09.  Fontanel absent i n a l l castes; present;  ocelli  t a r s i 5 - j o i n t e d i n a l l c a s t e s ; h i n d wings w i t h a w e l l  developed anal l o b e ; g e n i t a l i a w e l l developed* T h i s f a m i l y i s represented  by a p r i m i t i v e ,  northern A u s t r a l i a n s p e c i e s , Mastotermes d a r w i n i e n s i s The f o u r remaining  Froggatt.  f a m i l i e s have 4 - j o i n t e d or r a r e l y ,  i m p e r f e c t l y 5 - j o i n t e d t a r s i i n a l l c a s t e s , and hind wings without an a n a l lobe; g e n i t a l i a a r e l a c k i n g or v e s t i g i a l . 2. Hodotermitidae S j o s t e d t , '25. Fontanel  and o c e l l i absent" and pronotum  saddle-shaped and narrower than the head i n a l l c a s t e s ; a n t e r i o r wing s c a l e s s h o r t , not o v e r - l a p p i n g the p o s t e r i o r wing s c a l e s or wing  stubs. The Hodotermitidae a r e subterranean,  f e e d i n g t e r m i t e s of A s i a and A f r i c a , many r e s p e c t s these show a higher  grass-  Although p r i m i t i v e i n  s o c i a l development than the  Kalotermitidae. 3. K a l o t e r m i t i d a e S j o s t e d t , *25. Fontanel absent i n a l l c a s t e s ;  ocelli  u s u a l l y p r e s e n t ; pronotum f l a t , u s u a l l y broader than the head i n a l l c a s t e s ; a n t e r i o r wing s c a l e s l a r g e , o v e r - l a p p i n g the p o s t e r i o r wing s c a l e s .  No worker  caste.  In t h i s f a m i l y a r e grouped d r y - and damp-  i  -11wood termites of p o t e n t i a l economic importance of which Zootermopsis A n g u s t i c o l l i s and Z. nevadensis  are the B r i t i s h  Columbia r e p r e s e n t a t i v e s * 4, R h i n o t e r m i t i d a e L i g h t ,  '21.  F o n t a n e l p r e s e n t ; mandibles of s o - c a l l e d Leucotermes or S e r r i t e r m e s type (Holmgren*11); a n t e r i o r wing s c a l e l a r g e (except i n Psammotermea'); wing o f t e n r e t i c u l a t e and without  h a i r s ; pronotum of worker and  soldier f l a t ,  without  a n t e r i o r lobe* With the exception of Prorhinotermes a l l s p e c i e s are subterranean ance.  and of c o n s i d e r a b l e economic  R e t i c u l i t e r m e s hesperus i s the B.C.  import"  r e p r e s e n t a t i v e of  this family. 5. Termitidae L i g h t ,  '21.  Fontanel p r e s e n t ; mandibles never of Leucotermes or S e r r i t e r m e s type; a n t e r i o r wing s c a l e never l a r g e ; wing only s l i g h t l y r e t i c u l a t e i f at a l l ; wing membrane and wing margin more or l e s s h a i r y ; pronotum of s o l d i e r a'nc? worker narrow, w i t h u p l i f t e d median a n t e r i o r lobe* The T e r m i t i d a e are almost ical,  subterranean  exclusively trop-  forms which do not f e e d e x c l u s i v e l y  c e l l u l o s e and l a c k the protozoan  fauna which crowds the  a l i m e n t a r y t r a c t of other t e r m i t e s .  on  -12KEY  TO BRITISH COLUMBIA SPECIES.  /  ALATES.  1. Fontanel  absent; o c e l l i absent; no worker caste;  cerci  p l a i n l y 5- or 6 - j o i n t e d ; t a r s i 5»jointed ... Zootermopsis 2. Fontanel present;  o c e l l i present;  or 3 - j o i n t e d ; t a r s i 4-Jointed  a worker caste; c e r c i 2-  .. . . . R e t i c u l i t e r m e s hesperus.  2. D o r s a l s u r f a c e of body t y p i c a l l y brown; l e n g t h of wings u s u a l l y 23-25mm. o c c a s i o n a l l y s h o r t e r (very r a r e l y 20mm.), width of wings u s u a l l y g r e a t e r than 5mm.; a l l h a i r s on upper s u r f a c e of hind t i b i a e l e s s than one-half tioiae  diameter of  .»».««» ....«*•...».»•«.••»«.».»»»»»».  Q..^.  D o r s a l s u r f a c e of body t y p i c a l l y very dark brown; l e n g t h of wings about 17 mm. about 4 mm.;  (very r a r e l y 20 ram.), width, of wings  v a r i o u s h a i r s on upper s u r f a c e of h i n d  f u l l y one-half  diameter of t i b i a e  tibiae  ^....... nevadensis  SOLDIERS. 1. C e r c i prominent, of 3 or more j o i n t s ; t a r s i of 5 j o i n t s ; t i b i a with spines b e f o r e apex; no f o n t a n e l l e ; clypeus not b i l o b e d ; mandibles toothed but u n l i k e „».* Zootermopsis 2 C e r c i i n d i s t i n c t , of 1 or 2 j o i n t s ; t i b i a e at most with 2 - or 3 a p i c a l spines, t a r s i of not more than f o u r  joints  R e t i c u l i t e r m e s hesperus 2. P o s t e r i o r end of head p l a i n l y broader than the a n t e r i o r end, head index u s u a l l y about .82; head width-mandible  index  -13^ about .85 as  ( t y p i c a l l y l e s s than .9) very r a r e l y as  .94  Z.  high  angusticollis  Sides of head more n e a r l y p a r a l l e l , head index about head width-mandible index about 1.04 1.00)  very r a r e l y as low as  ( t y p i c a l l y more than  .98  Z. nevadensis  M o d i f i e d from "Bank's R e v i s i o n of N e a r c t i c Termites" Sumner's "The  .77,  and  Species of the Termite Genus Zootermopsis  Emerson". DISTRIBUTION OF THE BRITISH COLUMBIA SPECIES The  common damp-wood t e r m i t e , Zootermopsis  a n g u s t i c o l l i s , '^f he most n o r t h e r l y d i s t r i b u t e d wood-dwelling termite of North America! ( C a s t l e i n X o f o i d  '34), i s found i n  B r i t i s h Columbia (as f a r as c o l l e c t i o n s have been made) as f a r n o r t h as Skidegate !  3 l ) and  ( l a t . 53'N), Queen C h a r l o t t e Islands  i n the i n t e r i o r at Quesnel Lake ( l a t . 52'  (Banks '20).  (Beall  30'N)  On Vancouver I s l a n d the s p e c i e s occurs as f a r n o r t h  as Courtnay on the east s i d e and at Long Beach, T o f i n o on west c o a s t . River.  The  On  the mainland the n o r t h e r l y r e c o r d i s Powell  s p e c i e s occurs a l l a l o n g the Lower F r a s e r V a l l e y as  f a r east as Hope.  In the Gulf of Georgia  Texada I s l a n d ( l a t . 49* limit  the  40*).  i t has been found on  C a s t l e s t a t e s that i n i t s northern  i t reaches an a l t i t u d e of about 4000 f e e t and  i s the  s p e c i e s o c c u r r i n g i n the more humid c o a s t a l r e g i o n s . The  small damp-wood t e r m i t e , Z. nevadensis,  l e s s common and  the records are few  i n number.  is far  A l a t e s have been  -14taken as f a r n o r t h as Loughboro I n l e t  on  the  mainland and -on Vancouver I s l a n d from Nanaimo and Royal  Oak  near V i c t o r i a *  B e a l l claims that t h i s s p e c i e s i s the p r e v a i l i n g  one at V i c t o r i a . localities  ( l a t . 50' 40')  Two  a l a t e s and f o u r s o l d i e r s from v a r i o u s  ( a l l lowland) of Vancouver seem to t a l l y i n most  r e s p e c t s with Z. nevadensis.  Also certain s o l d i e r s i n a large  colony of Z. a n g u s t i c o i l i s have more or l e s s the measurements of Z. nevadensis.  As Sumner ('33)  characteristic states, hybrids  may  p o s s i b l y occur,  i n which case, an e x p l a n a t i o n may  for  such specimens which do not agree with the normal form.  s i n g l e a l a t e has been taken at Salmon Arm  (Leech).  be  The  found A  species  i s e s s e n t i a l l y a form found at higher a l t i t u d e s with a l s o a more n o r t h e r l y d i s t r i b u t i o n .  The r e c o r d f o r Quesnel Lake  may  be t h i s s p e c i e s * The western subterranean  termite R e t i c u l i t e r m e s  hesperus. occurs " s p a r i n g l y as f a r north as B r i t i s h states Pickens  ( K o f o i d '34)»  Columbia"  Records i n the U n i v e r s i t y c o l l e c t -  ion show that i t extends a l o n g the east coast of Vancouver I s l a n d i n areas having l e s s than t h i r t y inches of p r e c i p i t a t i o n per annum between V i c t o r i a and Departure Bay. where e l s e i n the c o a s t a l r e g i o n .  I t i s found  no-  In the i n t e r i o r i t i s d i s -  t r i b u t e d throughout the Okanagan V a l l e y from the border a s . f a r n o r t h as Vernon and a l s o at Kamloops and T r a n q u i l l e . New may  records  p l a c e i t i n the Upper F r a s e r Canyon above L y t t o n as w e l l .  -15FACTORS AFFECTING DISTRIBUTIONi Owing to the f a c t that there a r e not s u f f i c i e n t records i n B r i t i s h Columbia no d e f i n i t e statements may be made i n d i c a t i n g the dependence of d i s t r i b u t i o n on c l i m a t i c f a c t o r s . There a r e , n e v e r t h e l e s s ,  several interesting facts  the r e l a t i o n s h i p s of c l i m a t i c f a c t o r s and termite Humidity i s probably  the most important  concerning distribution.  of the c l i m a t i c f a c t o r s  a f f e c t i n g the range of Z. a n g u s t i c o i l i s . As the humidity the colony  within  i s n e a r l y 100% ( K o f o i d *34), i t i s e s s e n t i a l that  they l i v e i n a f a i r l y humid c l i m a t e .  Their d i s t r i b u t i o n i s  l i m i t e d to p a r t s of the p r o v i n c e having at l e a s t t h i r t y annual p r e c i p i t a t i o n .  inches  Consequently they a r e not found i n the  greater p a r t of the c e n t r a l i n t e r i o r . .East of here there i s no reason why t e r m i t e s should not be found, as i s evidenced one r e c o r d from Salmon Arm.  by the  I t i s of i n t e r e s t to note, however  that damp-wood t e r m i t e s have not been recorded from a l l p a r t s having a t o t a l p r e c i p i t a t i o n of t h i r t y p a r t s of the northern  inches, such as many  coast and c e r t a i n p a r t s of eastern  B r i t i s h Columbia, but f u r t h e r records may l a t e r extend  distrib-  u t i o n a l l i m i t s to these areas as w e l l . Temperature, almost i n s e p a r a b l e from humidity, affects distribution.  Of temperatures, the maxima and minima  are those of greatest importance and a r e probably f o r the northern  also  responsible  l i m i t on the coast.  As regards R e t i c u l i t e r m e s hesperus, humidity i s probably  the c h i e f f a c t o r a f f e c t i n g d i s t r i b u t i o n as the species  seems able to withstand extreme ranges of temperature. east coast of the I s l a n d i s found, the r a i n f a l l  On the  "between Nanaimo and V i c t o r i a where i t  i s l e s s than t h i r t y i n c h e s .  In the  i n t e r i o r where i t i s recorded, the annual p r e c i p i t a t i o n i s much • l e s s , averaging about  ten inches at both O l i v e r and  Tranquille.  It i s more common i n the southern Okanagan than f u r t h e r  north  where the p r e c i p i t a t i o n i s somewhat g r e a t e r . R, hesperus i s able to withstand lower a i r humidity because i t i s subterranean.  -17.EXTERNAL AEA.TOHY , ,  A Comparison  of the Three  Species„  THE HEAD. Alates Zootermopsis a n g u s t i c o l l i s The head i s prognathous and when viewed d o r s a l l y i s a somewhat f l a t t e n e d capsule, rounded behind, narrowed  i n front  and broadest immediately p o s t e r i o r to the compound eyes ( P l a t e 1, f i g . l ) .  The e p i c r a n i a l suture i s not d i s t i n c t i n  l i v e specimens but i n those cleared^i't appears as a Y-shaped l i n e , f o r k i n g at a p o i n t j u s t p o s t e r i o r to an imaginary l i n e j o i n i n g the p o s t e r i o r margins of the two compound eyes, each f o r k extending to the a n t e r o - d o r s a l margin of each eye. The f e n e s t r a e (Balfour-Browne '32) a r e s c a r c e l y d i s c e r n a b l e and have no connection w i t h the tentorium or any head muscles. The broad e l l i p t i c a l compound eyes a r e s i t u a t e d approximately i n the center of the l a t e r a l  s u r f a c e of the head about midway  between the d o r s a l and v e n t r a l s u r f a c e s . front  O c e l l i a r e absent*  The  i s f i v e - s i d e d , bare and bounded a n t e r i o r l y by the e p i -  stomal r i d g e i n t e r n a l l y and by the p i t s of the a n t e r i o r arms of the tentorium* over the genae.  L a t e r a l l y the f r o n s forms a prominent  ridge  The elongated pleurostomae of the subgenae are  c l e a r l y d e f i n e d on the l a t e r a l  sides of the head,  V e n t r a l l y the  hypostomae a r e p o o r l y d e f i n e d and a r e hidden by the m a x i l l a e . The postgenae, bounded l a t e r a l l y by very prominent  occipital  sutures, make up a l a r g e p a r t of the V e n t r a l surface of the head, the remainder b e i n g taken up by the postmentum.  The  -18clypeus i s stiort, i t s l e n g t h being about o n e - t h i r d i t s breadth. The p o s t c l y p e u s i s d i f f e r e n t i a t e d from the p r e - c l y p e u s i n being much darker i n c o l o u r .  The i n n e r surfjace of the labrum i s  continuous with the membranous inner s u r f a c e of the c l y p e a l r e g i o n , the two forming the epipharyngeal w a l l or epipharynx. This c o n s i s t s o f many slender setae p o i n t i n g m e d i a l l y on the right  side (Plate I I , f i g . 4).  Tormae i n the l a t e r a l  angles  between the labrum and the clypeus a r e v i s i b l e i n c l e a r e d specimens as slender s c l e r i t e s . left  In a l l specimens examined the  torma i s smaller and c o n s i s t s of two p a r t s s i t u a t e d at  r i g h t angles to each other.  The hypopharynx i s a more or l e s s  t r i a n g l e - s h a p e d lobe as seen l a t e r a l l y It extends  ( P l a t e I I , f i g s . 8 and 9 ) .  to the t i p s of the p a r a g l o s s a e .  The a d o r a l s u r f a c e of  the hypopharynx i s c l o t h e d with many slender setae which are most numerous at the proximal  end.  The hypopharynx i s continuous  through the mouth w i t h the v e n t r a l w a l l of the stomodaeum.  The  a d o r a l area i s f l a n k e d on each s i d e by a p a i r of s c l e r i t e s which make up the suspensoria. suspensorium  D i s t a f f and a r t i c u l a t i n g with each  i s a l a t e r a l s c l e r i t e which extends  to the s a l i v a r -  ium. Zootermopsis  nevadensis.  The head of t h i s species i s s l i g h t l y smaller than that of the former pect  s p e c i e s but i s l i k e i t i n every other r e s -  (Tables 1 and 2 ) .  -19Reticulitermes  hesperus.  The head i s prognathous, a more or l e s s capsule, "broadly e l l i p t i c a l as viewed d o r s a l l y  flattened  ( P l a t e 1, f i g . 2 ) .  The f e n e s t r a e above and between the compound eyes and antennae y .are prominent.  The  s i t u a t e d immediately  compound eyes are c i r c u l a r i n o u t l i n e , are p o s t e r i o r to the antennae and are  separ-  ated from them by an amount equal to the width of the compound eye* and  The l a t e r a l o c e l l i ,  l o c a t e d midway between the fenestrae  the compound eyes are s l i g h t l y a n t e r i o r to the latter„  median o c e l l u s i s l a c k i n g but i t s p o s i t i o n may  The  be occupied by  the f r o n t a l gland (Thompson 16) which i s l o c a t e d at the b i f u r 1  c a t i o n of the e p i c r a n i a l s u t u r e .  The p o s i t i o n i s i n d i c a t e d by  a l i g h t - c o l o r e d spot i n the c h i t i n .  It i s very u n l i k e l y i n the  a l a t e forms that there i s an opening  e x t e r i o r l y as there are  setae on the c h i t i n a t t h i s spot.  The  e p i c r a n i a l suture can  be seen as a f a i n t Y-ehaped l i n e b i f u r c a t i n g a l i t t l e behind a l i n e connecting the p o s t e r i o r margins of the compound Here i t becomes very f a i n t and d i s a p p e a r s . point  eyes.  A n t e r i o r to t h i s  the e p i c r a n i a l arms extend to the p o s t e r i o r margins of  the o c e l l i . although maxillae.  Both pleurostomae and hypostomae are p r e s e n t ,  the l a t t e r i s even more completely The o c c i p i t a l  suture i s not  hidden by  so apparent  but i t s  p o s i t i o n i s i n d i c a t e d by a r i d g e as i n Zootermopsis. i s much the same i n shape as that of Zooterraopsis.  the  The  The p o s t -  clypeus i s somewhat e l l i p t i c a l i n shape, two and one-half as broad as l o n g .  front  times  The preclypeus i s one-fourth the l e n g t h of  -20the p o s t c l y p e u s .  The labrum ( P l a t e I I , f i g . 5) has setae o f  two  s i z e s , the two c e n t r a l l a r g e ones "being very prominent.  The  setae making up the epipharynx  l i e c e n t r a l l y while i n  Zooterrnopsis they a r e i n a U-shaped f o r m a t i o n .  In a d d i t i o n  there i s a narrow row of l a r g e d e n t i c l e s without  spines  besides  many s m a l l e r ones i n R e t i c u l i t e r m e s while i n Zooterrnopsis the d e n t i c l e s , which are spined, a r e a l i k e i n s i z e and s c a t t e r e d evenly over  the whole s u r f a c e .  The tormae i n the l a t e r a l  angles  between the labrum and clypeus a r e t h r e a d - l i k e and somewhat curved.  The hypopharynx i s s u p e r f i c i a l l y the same as i n  Zooterrnopsis. Brachypteroua  Reproductives.  Zooterrnopsis a n g u s t i c o i l i s . The head i s s l i g h t l y broader  than that of the f i r s t  form a d u l t and tapers l e s s i n f r o n t of the eyes (Thompson '22). The  eyes a r e o v a l and not reniform, 0.2 by 0,1 mm.,  smaller than the eyes of f i r s t  slightly  form nymphs (Thompson '22).  In Table 3 i t w i l l be n o t i c e d that although the head i s wider than i n the a l a t e s i t i s much longer than broad, the r e v e r s e being t r u e f o r a l a t e s .  The head i s much p a l e r i n  c o l o r , only the clypeus and mandibles being dark.  The e p i c r a n -  i a l suture may be seen i n l i v e specimens as a s l i g h t Y-shaped depression of l i g h t e r c o l o u r . antennae a r e not as prominent.  The r i d g e s above the genae and The f e n e s t r a e are o v a l and  because the head i s l i g h t e r they are f a r more apparent. compound eyes may or may not be s l i g h t l y f a c e t e d .  The  In a l l other  -21r e s p e c t s t h i s caste  i s the same as the a l a t e s .  No specimens of t h i s caste f o r e i t h e r or R. hesperus were  Z. nevadensis  available.  Apterous R e p r o d u c t i v e s . Zooterrnopsis a n g u s t i c o i l i s . A c c o r d i n g to Thompson ('22) the compound eyes a r e of the same s i z e as those of the second form.  The head i s much  "broader. The head i s s l i g h t l y l a r g e r  (Tables 3 and 4 ) .  The  f e n e s t r a e , which here extend to the r i d g e above the antennae, are  s t i l l more n o t i c e a b l e as the head i s of a l i g h t c o l o u r .  The compound eyes a r e s l i g h t l y f a c e t e d . are  S c l e r i t e s and sutures  the same as i n the second form or brachypterous reproduc-  tives. No specimens of the t h i r d form or apterous reproduct i v e s of the other two s p e c i e s were a v a i l a b l e . Soldiers; Zooterrnopsis a n g u s t i c o l l i s . The head of s o l d i e r s i s very dark p a r t i c u l a r l y t o wards the mandibles and i s subject to much v a r i a t i o n i n s i z e . In the m a j o r i t y of cases the head i s about  one-fourth longer  than wide although i n other eases the head width equals the head l e n g t h (Table 5 ) . alates,  I t i s much more elongate than i n the  longer to accommodate the much l a r g e r  mandibular  -22muscles. although  ^Functional eyes are u s u a l l y absent i n t h i s  caste  i n c l e a r e d specimens a l i g h t e r - c o l o u r e d narrow o v a l  spot i s present "behind  each antennal  foveola.  In some i n d i v -  i d u a l s the eyes a r e broadly o v a l and a r e f u n c t i o n a l , as f a c e t s may be d i s t i n g u i s h e d . distinct heads.  i n l i v e s o l d i e r s but can be r e a d i l y seen i n c l e a r e d The p o i n t o f b i f u r c a t i o n of the e p i c r a n i a l suture i s  l o c a t e d approximately to an imaginary present  The e p i c r a n i a l suture and arms are not  midway but p o s t e r i o r , by a considerable,  l i n e j o i n i n g the eyes.  as spots t a p e r i n g l a t e r a l l y .  hypostomae a r e p r e s e n t . labrum i s longer  Both pleurostomae and  The o c c i p i t a l suture i s absent.  than broad (Table 5 ) .  ynx a r e a s y m e t r i c a l as  The f e n e s t r a e are  The  The tormae and epiphar-  i n the a l a t e s ( P l a t e I I , f i g . 6 ) .  26bteriaopais  nevadensis.  Although the head of the s o l d i e r of t h i s c l o s e l y resembles that of a n g u s t i c o l l i s i t d i f f e r s  species considerably  i n some of i t s measurements and i n d i c e s (Table 5 ) . The head i s n o t i c e a b l y much s m a l l e r .  The labrum i s emarginate d i s t a l l y  (Plate I I , f i g . 3). Ret i c u l l t ermes hesperus. S o l d i e r s as a r u l e i n d i c a t e to a marked degree the development of the s p e c i e s .  In comparing t h i s s p e c i e s with  gootermopsis as shown i n the p h y l o g e n e t i c  key ( P l a t e IV, £lg. l )  we f i n d that i t i s f u r t h e r advanced. The head v a r i e s from n e a r l y twice as l o n g to much  -23more than twice as l o n g as broad, b e i n g broadest at i t s ante r i o r margin and t a p e r i n g s l i g h t l y p o s t e r i o r l y . d u l l brownish-yellow  It i s of a  c o l o u r , much p a l e r than that of the other  B r i t i s h Columbia species*  The f o n t a n e l l e i s q u i t e d i s t i n c t ,  b e i n g l o c a t e d s l i g h t l y a n t e r i o r to the p o s t e r i o r margin of the brain.  A n t e r i o r to t h i s p o i n t the roof of the head has a marked  s w e l l i n g which i n v o l v e s most of i t s a n t e r i o r p o r t i o n ( P l a t e I I , f i g . 2).  T h i s s w e l l i n g i s only s l i g h t l y i f at a l l d i s c e r n a b l e  i n Zooterrnopsis ( P l a t e I I , f i g * r e l a t e d i n any way  l).  Whether the s w e l l i n g i s  with the b r a i n which l i e s immediately  or w i t h the f o n t a n e l l e i s hard to determine.  The  sutures are v i s i b l e only i n c l e a r e d specimens.  below  epicranial  Located on the  e p i c r a n i a l arms about two-thirds the d i s t a n c e (between the f o n t a n e l l e and the antennae) from  the f o n t a n e l l e i s a p a i r of  small l i g h t - c o l o u r e d c i r c u l a r s p o t s . imentary  ocelli.  These are probably rud-  L a t e r a l and v e n t r a l to each of these spots  and p o s t e r i o r to the lower margin of the antennal socket are two  other l i g h t - c o l o u r e d spots of the same s i z e , probably  r e p r e s e n t i n g compound eyes which are not f u n c t i o n a l . pleurostomae  does not form a d i s t i n c t  pointed d i s t a l l y  sclerite.  The  The labrum i s  ( P l a t e I I , f i g . 6) and i s longer than  broad.  Near i t s t i p are two prominent d i v e r g i n g setae while the whole surface i s covered w i t h many smaller setae.  The epipharynx i s  s l i g h t l y d i f f e r e n t i a t e d from the r e s t of the epipharyngeal w a l l .  -24THE COMPOUND EYES* 1  . Owing to the f a c t that the compound eyes  of  Zootermopsis a n g u s t i c o l l i s , and p o s s i b l y nevadensis as •well, are  subject to so much v a r i a t i o n , t h i s a d d i t i o n a l note i s  necessary*  In mature forms, other than a l a t e s and  soldiers,  that i s , brachypterous, apterous and 8th i n s t a r brachypterous, (which may  be the same as the f i r s t l i s t e d ) , have compound eyes  s i m i l a r to the 7th i n s t a r forms which w i l l moult alates*  to become  In these cases the eye as a whole i s convex, although  not to the same degree as i n the a l a t e s *  The cornea i s f a c e t e d  but only s l i g h t l y , being i n t e r m e d i a t e between the f a c e t a t i o n of  a l a t e s and the smooth c o n d i t i o n of normal  soldiers.  The  degree of pigmentation v a r i e s s l i g h t l y - i n most apterous.forms it  i s s l i g h t w h i l e i n others i t i s p a l e to dark brown.  One  apterous i n d i v i d u a l had f a c e t e d eyes without any pigment (an a l c o hol  specimen).  The pigmentation v a r i e s i n the eye  itself,  b e i n g darker i n the v e n t r a l p o r t i o n than i n the d o r s a l . eyes i n a l l of these i s about nymphs. pigmented  The  the same s i z e as that of 7th i n s t a r  In one apterous form the eye was  smaller, s l i g h t l y  and not f a c e t e d . One  s o l d i e r examined had pigmented  eyes with i n t e r -  mediate f a c e t a t i o n .  In t h i s specimen  s l i g h t l y pigmented.  In a l l the other s o l d i e r s , the eye surface  i s smooth and i s n e a r l y f l a t .  the eye was  convex and  The eyes are of the same c o l o u r  as the r e s t of the head and are smaller than i n the other castes.  -25Th e nearest approach to the compound eyes of the alates* i s found i n the 8th i n s t a r brachypterous r e p r o d u c t i v e s i n which pigmentation  i s dark brown.  Immature stages; Zooterrnopsis a n g u s t i c o i l i s In f i r s t and  i n s t a r nymphs the head i s broader  than l o n g  i s densely c l o t h e d with l o n g slender setae ( P l a t e 1, f i g .  5 and 6 ) . arms.  There i s no t r a c e of the e p i c r a n i a l suture or i t s  Eyes are e n t i r e l y absent.  indistinct. alates.  The epistomal suture i s very  The f r o n t forms a s l i g h t r i d g e over the genae as i n  The o c c i p i t a l sutures are not p r e s e n t .  not d i f f e r e n t i a t e d i n t o i t s two p a r t s *  The clypeus i s  The epipharyngeal w a l l  i s only s l i g h t l y d i f f e r e n t i a t e d i n t o an epipharynx, f i f t e e n short but  stout setae on the r i g h t s i d e .  not yet become s c l e r o t i z e d . as that of a l a t e s and extends Other  there being  Tormae have  The hypopharynx i s the same shape s l i g h t l y beyond the paraglossae.  s c l e r i t e s mentioned i n connection w i t h the a l a t e s have  not become d i f f e r e n t i a t e d . In t h i r d i n s t a r nymphs the head has much fewer setae\ which are s m a l l ,  The e p i c r a n i a l suture and arms are l a c k i n g  while the epistomal suture i s very s l i g h t l y marked.  The  pharynx and tormae are much the same as i n the a l a t e s but left  epithe  tormae c o n s i s t s of only one p i e c e , the small median  s c l e r i t e i n connection with i t i n the a l a t e s i s l a c k i n g .  The  hypopharynx extends n e a r l y to the t i p s of the l a c i n a e but not as f a r as the t i p s of the paraglossae*  -26Immature stages of the other s p e c i e s were not a v a i l able. ' THE  ANTEMAE. Alates Zootermopsis  angusticollis  The antennae are l o n g and s l i g h t l y more than twice the  g r e a t e s t width of the head^aadrthe  segments m o n i l i f o r m .  The mean number of antennal segments i s t w e n t y - f i v e with a range of between twenty-four and twenty-seven.  The scape i s  s l i g h t l y l o n g e r and c o n s i d e r a b l y broader than the next segments.  two  These l a t t e r are n e a r l y quadrate i n o u t l i n e *  The  f o u r t h segment, the s m a l l e s t , i s twice as broad as i t i s long. Distally  the segments become s l i g h t l y broader and longer |to the  most d i s t a l , which i s twice as l o n g as i t i s broad.  This  segment and a few immediately p r e c e d i n g are p e t i o l a t e d . Zootermopsis nevadensis The antenna^ of t h i s s p e c i e s i s l i k e that of the preceding  species. R e t i c u l i t e r m e s hesperus The antennae o f t h i s  species are twice as l o n g as the  head and c o n s i s t of seventeen m o n i l i f o r m segments. examined had eighteen*.  The t h i r d  One  specimen  segment i s the smallest  a l l others distal^f/become i n c r e a s i n g l y l o n g e r , the terminal b e i n g twice as l o n g as broad.  and ®VJS  As i n Zootermopsis the scape i s  as l o n g a a ^ and s l i g h t l y b r o a d e r ^ than^the next two  segments.  Brachypterous '  Reproductives  Zooterrnopsis a n g u s t i c o i l i s . In three i n d i v i d u a l s examined the antennae c o n s i s t e d  of twenty-four,  twenty-five and  twenty-six  segments.  The  scape  iVian  ±3 much "broader and as l o n g as ,,the next  two  segments combined.  The f o u r t h segment i s very small b e i n g one f o u r t h as long as broad.  Segments f i v e to ten are a l l broader  than l o n g , while  the remainder become g r a d u a l l y longer to the d i s t a l segment which i s ' s l i g h t l y s m a l l e r .Ytam "Hie penuftvmate. No  specimens of brachypterous  f o r examination-,,in? the other two Apterous  forms were a v a i l a b l e  species^  Reproductives.  Zooterrnopsis a n g u s t i c o i l i s * The antennal  count f o r t h i s caste v a r i e s considerably  but u s u a l l y i t i s about twenty-five or s i x . twenty-nine*  specimen had  The f o u r t h and f i f t h segments are the s m a l l e s t .  No apterous two  One  r e p r o d u c t i v e s were obtained f o r the  other  species. Soldiers* Zooterrnopsis angus.t i c o l l i s . The ant e n n a ^ t h i s  number of antennal  segments^ depending on the i n s t a r i n which  the s o l d i e r bectwneSmature. the v a r i a t i o n was of t w e n t y - f i v e .  case v a r i e s somewhat i n the  Of counts of twenty-four  eighteen to twenty-eight The  soldiers,  with a mean number  f o u r t h segment i s the s m a l l e s t .  -28Zooterrnopsis nevadensis ' The number of antennal j o i n t s v a r i e s from twenty to twenty-three or twenty-four (Heath and Wilbur  '27).  R e t i c u l i t e r m e s hesperus. The antennae of t h i s caste c o n s i s t of f i f t e e n  segments.  Immature Stages. Zooterrnopsis a n g u s t i c o i l i s The newly hatched nymph i n the f i r s t antennae  o f f o u r t e e n segments.  i n s t a r possesses  The second i n s t a r has  sixteen  antennal j o i n t s ; the t h i r d , eighteen; the f o u r t h twenty; f i f t h u s u a l l y twenty-two but sometimes twenty or the  the  twenty-one;  seventh u s u a l l y t w e n t y - f i v e . As i n the s l a t e s the f o u r t h  segment i s the smallest* Zooterrnopsis nevadensis. The newly hatched t e r m i t e has twelve or t h i r t e e n segments to each antenna.  The second has f i f t e e n ; the t h i r d ,  seventeen; the f o u r t h , eighteen or n i n e t e e n ; the f i f t h , nineteen to  twenty-one (Heath and Wilbur '27)* R e t i c u l i t e r m e s hesperus A s e r i e s of the immature stages of t h i s s p e c i e s was  not  available.  -29THE MOUTHPARTS. THE MANDIBLES. Alates Zootermopsis a n g u s t i c o l l i s Prom e i t h e r v e n t r a l or d o r s a l aspect,  the mandibles  are quadrate i n o u t l i n e and have a t r i a n g u l a r base. from the t i p of the condyle to the d i s t a l shorter than the gulamentum.  end they a r e s l i g h t s  The dentes a r e very dark brown to  b l a c k , the mandible becoming straw-coloured The  Measuring  towards i t s base.  r i g h t mandible has four t e e t h i n which the pen-  u l t i m a t e i s the smallest  ( P l a t e I I I , f i g , 1A and B).  The l e f t  mandible has a l s o f o u r t e e t h b u t a r e s l i g h t l y smaller and of A  different  shape and p o s i t i o n .  The mola of the r i g h t mandible i s  c r e n u l a t e f o r i t s e n t i r e l e n g t h , s l i g h t l y curved one-half  and i s n e a r l y  the l e n g t h of the inner surface of the mandible.  The  mola of the l e f t mandible i s smaller, of i r r e g u l a r shape and only very  slightly  crenulate.  (counting from the l e f t  A r i s i n g from the f o u r t h r i d g e  v e n t r a l side) of the mola of the l e f t  mandible there i s a p l a t e - l i k e structure-.  The c r e n u l a t i o n s of  the mola a r e not as long i n the r i g h t mandible as i n the l e f t , numbering u s u a l l y f i f t e e n on the r i g h t and twelve on the l e f t . Except f o r very minute setae,  the scrobe i s bare.  tendon i s smaller than the recto-tendon  The extenso-  and s h o r t e r than thefwhae  mandible. Zootermopsis nevadensis The mandibles of t h i s species a r e not d i s t i n g u i s h a b l e  -30from those of the p r e c e d i n g  species,  R e t i c u l i t e r m e s hesperus. The mandibles of t h i s species d i f f e r i n that penultimate d i s t a d e n t i s of the l e f t mandible i s smaller the ante-penultimate.  The proxadentis  ( P l a t e I I I , f i g . 2A and  the than  i s relatively larger  B).  Brachypterous and Apterous  Reproductives.  Mandibles of these castes do not d i f f e r from those of the a l a t e s of Z.  angusticoilis.  Soldiers. Zooterrnopsis  angusticoilis.  The mandibles here are three times as long as a l a t e mandible ( P l a t e I I I , f i g . 3A and B ) . measurements and  In t a b l e 5 are  i n d i c e s f o r purposes of comparison.  a l a t e s the r i g h t and l e f t mandible has  tooth the A  given  As i n the  l e f t mandibles are s l i g h t l y d i f f e r e n t .  f o u r t e e t h of which the second i s the  (measured along an a n t e r o - p o s t e r i o r l i n e ) , and proximal  the  shortest.  The  longest  the f o u r t h and  P o s t e r i o r to the l a t t e r i s the  mola which c o n s i s t s of a t o o t h with two  distinct projections  and  c o n s i s t ^ of a s e r i e s of seven c r e n u l a t i o n s v e n t r a l and p o s t e r i o r to the t o o t h .  Of the r i g h t mandible the most d i s t a l of the  teeth i s the l o n g e s t , the others are of equal has d e n t a t i o n s  The mola  over i t s e n t i r e surface, numbering u s u a l l y  twenty-one, the p o s t e r i o r being four-tenths  length.  three  smalle^.  the l e n g t h of the mandible and  The  extenso-tendon i s  o n e - f i f t h as broad  -31as l o n g .  The recto-tendon  is slightly  mandible"and c o n s i s t s of a broad f l a t  shorter than the  left  (nearly h o r i z o n t a l )  v e n t r a l p l a t e , o n e - t h i r d as wide as l o n g , u n i t e d a n t e r i o r l y with a narrower o b l i q u e p l a t e .  The h o r i z o n t a l p l a t e i s strengthened  by a t h i c k e n i n g of the i n n e r l a t e r a l margin (and') at the p o i n t where i t i s u n i t e d with the mandible. Zootermopsis nevadensis. The mandibles of t h i s species are longer narrower than those of an^us^coJJ.i_s_iTable 5^ ^ P l a t e I I I , f i g . 5A and  and  and  B).  R e t i c u l i t e r m e s hesperus The mandibles of the s o l d i e r caste bear  little  resemblance to those of a l a t e s ( P l a t e I I I , f i g s . 4A and In l e n g t h , they are s l i g h t l y shorter than the width of head and are s h a r p l y p o i n t e d meson.  The  The recto-tendon  the  or three .  the mola i s the most prominent.  of the l e f t mandible i s compound as i n the  s o l d i e r s of Zooterrnopsis and extending  the  curving much d i s t a l l y toward  teeth on the mandible are reduced to two  small p r o j e c t i o n s ; of'these  B).  i s as l o n g as the mandible  itself,  p o s t e r i o r l y to the o c c i p i t a l foramen. According  to the characters of the mandibles of  s o l d i e r s a phylogenetic  the  key has been prepared ( P l a t e IV, f i g . l ) .  (Hare ^ i ) .  Zooterrnopsis  angusticollis.  The mandibles of the f i r s t  i n B t a r are  superficially  quite d i f f e r e n t from those of the a l a t e s ( P l a t e I I I , f i g s .  -326A and B), but on c l o s e r examination i t w i l l be found that number' of i n d e n t a t i o n s and p r o j e c t i o n s i s the same. the t e e t h are not at a l l prominent and  the  However,  the r i d g e s on the mola  are s c a r c e l y i f at a l l p r e s e n t . The mandibles of the t h i r d i n s t a r have the proxad e n t i s of the l e f t  mandible very acute a p i c a l l y , and  ridgeB  of the mola more prominent than i n a l a t e s . THE  MAXILLAE Alates Zootermopsis a n g u s t i c o l l i s The m a x i l l a e are more or l e s s elongated  figi  1 and  2).  proximally.  The  The  tapering  stipes i s rectangular,being  l o n g as broad, being d i s t a l end  cardo i s r e c t a n g u l a r  ( P l a t e V, slightly  n e a r l y twice  s l i g h t l y f l a r e d and undulated at  to r e c e i v e the g a l e a and l a c i n a .  The  as  the  subgalea,  only s l i g h t l y d i f f e r e n t i a t e d from the r e s t of the galea, i s roughly tip  c i r c u l a r and  o n e - t h i r d the width of the s t i p e s .  The  of the galea i s c l o t h e d with numerous short setae p o i n t i n g  inwards and lacina.  The  f o l d s h o o d - l i k e around the d i s t a l p o r t i o n of  l a c i n a i s h a l f again as l o n g as the galea and i s  equipped at the t i p with two l a t e r a l and  l a r g e acute t e e t h and  v e n t r a l surfaces with stout setae.  i s composed of eighteen  very l a r g e setae.  The  the palpus i s n e a r l y as l o n g as the f i r s t  two.  fifth longer  the  on  the  The l a c i n a r a s t r a t h i r d segment of The  fourth  and  segments are of n e a r l y equal l e n g t h , each about o n e - t h i r d than the t h i r d segment.  The  distal joint  i s narrower  -33-; than any  of the o t h e r s .  A l l f i v e p a r t s of the palpus have Ion  setae about as long as the width of the d i s t a l segment. a d d i t i o n to these there are a number of much smaller  In  setae.  With the exception of the setae of the d i s t a l segment, a l l are l o c a t e d on the median two-thirds of the segment.  Those of the  d i s t a l ^ e x t e n d from c l o s e to the base, to the end of the segmenZooterrnopsis  nevadensis  The m a x i l l a e are s l i g h t l y s m a l l e r .  In the specimens  examined the l a c i n a r a s t r a u s u a l l y c o n s i s t of seventeen stout setae, although  i n one there were eleven,  In other r e s p e c t s  they are i d e n t i c a l with those of a n g u s t i c o i l i s . R e t i c u l i t e r m e s hesperus. There are s e v e r a l morphological  characters d i s t i n g -  u i s h i n g the m a x i l l a e of t h i s species from the two s p e c i e s ( P l a t e V, f i g . 3 and 4 ) . of eleven l a r g e setae.  The  The l a c i n a r a s t r a i s composed  galea i s much l a r g e r ,  w e l l above the t e e t h of the l a c i n a and i t s l e n g t h towards the meson.  preceding  The  is slightly  extending curved i n  t e e t h of the l a c i n a are  correspondingly longer than i n Zootermqpsis.  The  setae on  v e n t r a l surface of the l a c i n a are much fewer, but l o n g e r . subgalea  the The  is distinct. Brachypterous  and Apterous  Reproductives.  In Z. a n g u s t i c o i l i s , the m a x i l l a e of these are s i m i l a r to those of a l a t e s .  castes  Soldiers •  Zootermopsis a n g u s t i c o l l i a . The m a x i l l a e o f t h i s caste a r e more h e a v i l y s c l e r o t -  ized.  Here the t h i r d segment of the m a x i l l a r y palpus i s longer  .than the "basal two and the f i f t h  segment i s of the same l e n g t h  as the "basal and equal to the combined l e n g t h s o f the second and t h i r d .  Setae of a l e n g t h n e a r l y equal to the width of the  segments are l o c a t e d on the three d i s t a l segments.  A few and  very small setae a r e present  on the "basal two *  equal i n l e n g t h to the f i r s t  three segments of the m a x i l l a r y  palpi.  The galea i s  The l a c i n a r a s t r a i s composed of nineteen Zootermopsis  setae.  nevadensis  In the one specimen examined, two f i f t h s of the galea extended above the teeth of the l a c i n a . c o n s i s t e d of f i f t e e n  setae.  The l a c i n a r a s t r a  In other c h a r a c t e r s , the m a x i l l a e  resemble those of a n g u s t i c o l l i s . R e t i c u l i t e r m e B hesperus The m a x i l l a e are s u p e r f i c i a l l y the same as those of the a l a t e .  There i s t h i s d i f f e r e n c e , however, each segment of  the m a x i l l a r y p a l p i a r e longer than the corresponding of  the a l a t e s , the proximal  segments  segments b e i n g more elongate*  The  l a c i n a r a s t r a c o n s i s t s of eleven setae, although one specimen had only n i n e .  Immature Stages * )  2oot ermopsis a h g u a t i c o l l i s In the f i r s t  i n s t a r , the l a c i n i a of the m a x i l l a  has at i t s t i p two very small and very acute t e e t h . l a c i n a r a s t r a c o n s i s t s of three short and  The  stout setae*  In the t h i r d i n s t a r examined, the d i s t a l teeth of the l a c i n a are l a r g e as i n the a l a t e s , and the l a c i n a r a s t r a has  twelve l a r g e setae of unequal length*  THE .LABIUM Alates Zootermopsis a n g u s t i c o l i i s The labium c o n s i s t s of a broad prementum and i t s two p a l p i g e r s and f o u r unsegmented glossae d i s t a l l y and a post mentum p r o x i m a l l y  ( P l a t e I, f i g *  2).  The prementum i s s l i g h t l y  more than twice as broad as l o n g , the l a t e r a l margins ing  proximally.  maxillary p a l p i .  The p a l p i are l e s s than h a l f the l e n g t h of the The  d i s t a l segment of each palpus  the l e n g t h of the f i r s t  two  combined*  i s equal to  The paraglossae are blunt  and almost meet above the glossae which are c o n i c a l and pointed*  The p a r a g l o s s a e are c l o t h e d with f a i r l y  while the glossae have only a few mesarima extends p o s t e r i o r l y one tum. two  converg-  very  sharply  l a r g e setae  small b r i s t l e s *  The  t h i r d the width of the premen-  The postmentum i s h e a v i l y s c l e r o t i z e d f o r i t s p o s t e r i o r t h i r d s and i s s p a r s e l y c l o t h e d with small setae s c a t t e r e d  over i t s surface* posteriorly.  It tapers to l e s s than one h a l f i t s width  -36Zootermopsis ;  except  nevadensis  The labium of t h i s s p e c i e s i s the same as the above  that the postmentum tapers l e s s p o s t e r i o r l y . R e t i c u l i t e r m e s hesperus In t h i s s p e c i e s , the prementum i s longer than i n  a n g u s t i c o i l i s ( P l a t e I, f i g . 4 ) . The postmentum i s j u s t a l i t t longer than the prementum. Brachypterous  and Apterous  Reproductives  The labium i s the same as i n the a l a t e s . Soldiers Zooterrnopsis a n g u s t i c o i l i s In the labium of t h i s caste the l a b i a l p a l p i a r e equal i n l e n g t h to the combined l e n g t h s o f the t h i r d and f o u r t h segments of the m a x i l l a r y p a l p i  ( P l a t e VI, f i g . l ) . The  gulamentum makes up the l a r g e r area o f the v e n t r a l s i d e of the head.  I t s v a r i o u s s i z e p r o p o r t i o n s a r e i n d i c a t e d i n Table 5.  The a n t e r i o r t i p of the gulamentum i s much p a l e r and i s the broadest  p a r t of the s t r u c t u r e .  It tapers p o s t e r i o r l y to a  p o i n t f o u r - f i f t h s the l e n g t h where i t broadens as i t approaches the o c c i p i t a l foramen.  In some i n d i v i d u a l s the gulamentum i s  p r a c t i c a l l y devoid of any setae. Zootermopsis  nevadensis  The labium o f t h i s s p e c i e s i s much the same as i n angustlcQ.11 i s .  Measurements of the gulamentum a r e given i n  -37Table 5. R e t i c u l i t e r r a e s hesperus The  l a b i a l p a l p i although of the same l e n g t h are  s l i g h t l y more slender  (Plate VI, f i g . 2 ) .  The gulamentum i s  f u l l y two and one h a l f times as broad i n f r o n t as at i t s narrowest  part.  THE TENTORIUM Alates Zootermopsis a n g u s t i c o l i i s As  i n other i n s e c t s the tentorium c o n s i s t s of (lj  the a n t e r i o r arms i n v a g i n a t e d  at .the f r o n t o - c l y p e a l  suture  j u s t mesad of the p r e c o i l a ; and (2) the p o s t e r i o r arms  invaginated  at the p o s t e r i o r l a t e r a l borders of the o c c i p i t a l foramen p a s s i n g forward to meet the a n t e r i o r arms^thus forming the corpotentorium. In t h i s species arms at a p o i n t  the p o s t e r i o r arms meet the a n t e r i o r  immediately v e n t r a p to the b i f u r c a t i o n of the  e p i c r a n i a l suture ( P l a t e VI, f i g . 4 ) . A p a i r of median processes a r i s i n g at the bases of the a n t e r i o r arms u n i t e  i n an a n t e r i o r  b r i d g e before the circumoesophageal connectives. arms a r e broadest at t h e i r p o i n t s  of i n v a g i n a t i o n .  p o i n t midway along t h e i r l e n g t h a r i s e two t a p e r i n g Their i d e n t i t y i s u n c e r t a i n arms.  but p o s s i b l y  The a n t e r i o r From a processes.  they are the d o r s a l  There are two tendons on the p o s t e r i o r t i p s of the  p o s t e r i o r arms: the d o r s a l apodeme and the tendon of the extensors of the head.  The tentorium o f 2„ nevadensis  i s l i k e that of the  preceding* s p e c i e s . Reticulitermes  hesperus  There a r e many marked d i f f e r e n c e s i n the tentorium of t h i s s p e c i e s ( P l a t e V I , f i g . 3 ) , The a n t e r i o r arms form a much broader "bridge a n t e r i o r to the circumoesophageal ives.  connect-  T h i s b r i d g e together with the p a r t of the corpotentorium  j u s t p o s t e r i o r to the neuroforamen i s very n e a r l y f l a t .  At the  body o f the tentorium the p o s t e r i o r arms a r e c o n s i d e r a b l y more separated. Brachypterous  and Apterous  Reproductives  Zooterrnopsis a n g u s t i c o i l i s Except f o r being r e l a t i v e l y l a r g e r , the t e n t o r i a of these c a s t e s i s the same as i n the a l a t e s ( P l a t e V I I , f i g , l ) . T e n t o r i a of the other two species were not examined. Soldiers Zooterrnopsis a n g u s t i c o i l i s The tentorium of t h i s caste i s very much modified, b e i n g very long with the b a a a l p a r t or corpotentorium a r o o f over the v e n t r a l nerve  cord" !and A  forming  the s a l i v a r y ducts and  extending from a p o i n t s l i g h t l y a n t e r i o r to the p o s t e r i o r margin of the gulamentum to an imaginary l i n e drawn between the eyes ( P l a t e V I , f i g . 5 ) . At t h i s p o i n t i t gives o f f v e n t r a l i y two short and stout p r o j e c t i o n s which extend to the a r t i c u l a t i o n of the m a x i l l a e with the head.  Prom each of these p r o j e c t i o n s  -39a r i s e s an a n t e r i o r extension -which g i v e s support and l a t e r a l p a r t of the gulamentum.  to the d i s t a l  The a n t e r i o r arms are very  "broad d o r s o - v e n t r a l l y at t h e i r p o i n t s of i n v a g i n a t i o n .  Dorsal  arms, as very t h i n p l a t e s t a p e r i n g p o s t e r i o r l y , a r i s e from the a n t e r i o r arms above the corpotentorium.  The neuraforamen i s  the same s i z e (by a c t u a l measurement) as i n the a l a t e s although i t appears smaller by c o n t r a s t with the r e s t of the Zoot ermopsis  tentorium.  nevadensis  In t h i s s p e c i e s the tentorium i s the, same as i n the preceding. R e t i c u l i t e r m e s hesperus In t h i s species as i n a n g u s t i c o l l i s the torium i s much elongated are very prominent.  ( P l a t e VI, f i g . 6 ) .  corpoten-  The a n t e r i o r arms  The b r i d g e or l a m i n i t e n t o r i u m and  the ape'  corpotentorium very narrow.  immediately  p o s t e r i o r to the neuraforamen  -jrS  3  In t h i s , the very small neura'formamen i s about  c e n t r a l l y placed*  The whole of the corpotentorium as i n the  a l a t e s a s q u i t e f l a t ; "Here too, there i s an a n t e r i o r p r o j e c t i o n on each s i d e of the corpotentorium,  g i v i n g support  to the  d i s t a l p a r t of the gulamentum. Immature Stages Zootermopsis a n g u s t i c o l i i s The t e n t o r i a of f i r s t  i n s t a r s are f l a t t e r and more  simple than i n a l a t e s ( P l a t e V I I , f i g . 2 ) .  The  corpotentorium  i3 p l a c e d more p o s t e r i o r l y i n r e l a t i o n to the head capsule,  -40making the former somewhat smaller, and the p o s t e r i o r arms s h o r t e r . ' The a n t e r i o r arms a r e r e l a t i v e l y longer and i n s t e a d of extending almost l a t e r a l l y as they do i n a l a t e s ^ t h e y nearly-anteriorly. larger.  extend  The neuraforamen i s p r o p o r t i o n a t e l y much  At the'-.posterior>tips,  the p o s t e r i o r arms a r e almost  parallel. In the second instar-j development i s towards that of the a l a t e tentorium more oblique,  ( P l a t e V I I , f i g . 3) w i t h the a n t e r i o r arms  the movement of the r e l a t i v e l y l a r g e neuraforamen  a n t e r i o r l y and an increase i n the s i z e of the^.-tentorium. As with a l l the-other •T  s t r u c t u r e s of the head of the  I  . alsc. ... 1  t h i r d i n s t a r , t h e tentorium feo#fc i s almost i d e n t i c a l with that of a l a t e s .  The a n t e r i o r arms a r e q u i t e o b l i q u e and the neura-  foramen i s n o t i c e a b l y r e l a t i v e l y smaller i n the younger  i n proportion,^ than  stages.  In c o n c l u s i o n  i t i s i n t e r e s t i n g to note t h a t ,  although the r a t i o of the head width of a l a t e s to the head width of newly hatched nymphs i s approximately 3.5:1, the r a t i o .of the s i z e o f the neuraforamen of the two stages i s only 1.32:1.  In examination of an e x u v i a S i t was seen that the t e n -  torium was shed at the same time as the r e s t of the  exoskeleton.  -41THJS CERVIX Alates Zootermopsis a n g u s t i c o l i i s and Z. nevadensis The  c e r v i x i s that p a r t  0  of the i n s e c t between the  "head and thorax and c o n s i s t s of .a c e r v i c a l membrane i n which ikese  there a r e f i v e s c l e r i t e s i nfettansspecies and  2).  ( P l a t e V I I I , figC 1  There i s a median v e n t r a l s c l e r i t e s i t u a t e d  p o s t e r i o r to the postmentum. on the l a t e r a l  immediately  P o i n t i n g v e n t r a l l y and extending  sides of the cervicum i s a p a i r of a n t e r i o r  lateral sclerites.  Extending toward the meson and s l i g h t l y  c a u d a d i s the p o s t e r i o r l a t e r a l 0  cervical scleritei'. /*  */ Reticulitermes The  hesperus.  c e r v i x i s l i k e that  w i t h the exception that  of the above two  species  the median s c l e r i t e i s l a c k i n g .  Brachypterous and Apterous Reproductives. In these castes i d e n t i c a l with that  i n Zootermopsis sp. the c e r v i x i s  of the a l a t e s .  Soldiers Zootermopsis a n g u s t i c o l i i s . The  c e r v i x i s s i m i l a r to that o c c u r r i n g  a l a t e s except that  the median s c l e r i t e may be l a c k i n g and that,  the a n t e r i o r l a t e r a l sclerotized.  c e r v i c a l s c l e r i t e s ' a r e much more h e a v i l y  In some i n d i v i d u a l s the median s c l e r i t e was  r e p r e s e n t e d by two very small of the meson.  i n the  s c l e r i t e s , placed  In others i t was complete, that  on each side i s of the same  -42s i z e as i n the a l a t e s .  The a n t e r i o r l a t e r a l  i n a d d i t i o n to b e i n g more h e a v i l y and  cervical sclerite  s c l e r o t i z e d i s also  shorter  3touter. Zooterrnopsis  nevadensis«  In the one specimen examined was somewhat longer*  In other r e s p e c t s  the median  sclerite  the c e r v i x i s the same  as i n a n g u s t i c o i l i s . R e t i c i a l i t e r m e s hesperus. The c e r v i x c l o s e l y resembles that anterior l a t e r a l s c l e r i t e s are s l i g h t l y  of a l a t e s .  The  longer.  THE THORAX. THE NOTA Alat e Zooterrnopsis a n g u s t i c o i l i s . Plates VIII, dorsal surface. twice i t s l e n g t h .  IX and X show the s c l e r i t e s of the  The pronotum width i s s l i g h t l y more than There i s a w e l l marked pleuranotum  (Plate  V I I I , f i g . l ) separated from the d o r s a l p a r t by a prominent ridge.  A n t e r i o r l y the pronotum  f i g s . 3 and 4 ) .  turns v e n t r a l l y ( P l a t e XI,  The median suture i s quite d i s t i n c t a n t e r i o r l y  as a l i g h t - c o l o u r e d l i n e which becomes f a i n t p o s t e r i o r l y . There a r e two quite dark areas near the a n t e r i o r margin of the pronotum, midway between the median suture and the r i d g e above mentioned.  These spots may be the p o i n t s of i n v a g i n a t i o n  -43of the endoskeleton  of the pronotum to "be d e s c r i b e d l a t e r ^  ija-^ii^-ip-a^ess-sThe mesonoturn ( P l a t e IX, f i g s . 3 and 4) h a s s e v e r a l distinct  areas.  A n t e r i o r l y the praescutum forms the preph-  ragma (Plat e XII, f i g . LA).  F o l l o w i n g t h i s i s the scutum and  the s c u t e l l u m with i t s l a t e r a l p o r t i o n s . wing process i s more c l e a r l y i o r notal-wing p r o c e s s . three a x i l l a r y sigmoidea  The a n t e r i o r n o t a l -  d i f f e r e n t i a t e d than the p o s t e r -  As i n most winged i n s e c t s , there are  sclerites.  The f i r s t M a x i l l a r y  i s eresent-shaped,  s c l e r i t e > or  the second or submedia i s some-  what o v a l i n shape and the t h i r d or t e r m i n a l i a i s t r i a n g u l a r and c o n s i s t s of two p a r t s .  In a d d i t i o n to these  sclerites  th^-r-e-is a f o u r t h ^ a r t i c u l a t i n g with the d i s t a l margin of the t h i r d a x i l l a r y on the one s i d e and connected with the wing on the o t h e r .  T h i s i s the jugum (Kalshoven  A n t e r i o r to the a x i l l a r i e s and connected  i n Sumner  '33).  with the c o s t a l  margin of the wing i s a small convex s e t i f er,ousf p a d - l i k e area, the t e g u l a . axillary  At the p o s t e r i o r marginals  s i t u a t e d the  cord which extends along the caudal margin of each  l a t e r a l p o r t i o n of the s c u t e l l u m .  The d o r s a l suture i s very  prominent as a dark l i n e i n t e r n a l l y  marked "by a p l a t e  des-  c r i b e d under exoskeleton. The metanotum i s somewhat short er and broader  than  the mesonotum ( P l a t e X, f i g . 1 ) .  The t e g u l a i s about  h a l f the 3 i z e of the mesotegula.  The jugum,-is not p r e s e n t .  The a x i l l a r y  W  s c l e r i t e s are s m a l l e r than i n mesonotum.  one-  T h i s d e s c r i p t i o n a p p l i e s to Zooterrnopsis nevadensis  except  that the p a r t s are s l i g h t l y smaller* Reticulitermes  hesperus.  The nota i n t h i s species are i l l u s t r a t e d i n "Plate X, f i g , 2,3,  and 4.  The pronotum i s p r o p o r t i o n a t e l y  much longer than i n the p r e c e d i n g species ( P l a t e IX, f i g . 2 ( 1 and 2 ) , portion.  The pleuranotum i s d i f f e r e n t i a t e d from the d o r s a l The a n t e r i o r p o r t i o n i s "bent v e n t r a l l y .  The d o r s a l  suture i s a l i g h t - c o l o u r e d l i n e prominent a n t e r i o r l y  and  posteriorly.  pre-  l\To dark spots are evident as i n the two  ceding s p e c i e s . The mesonotum ( P l a t e X, f i g s . 2 and 3) i s much the same as i n Z0otermopsis but the r e s t of the notum.  the scutum i s much shorter than  The other areas of the noturn are  d i s t i n g u i s h a b l e as the praescutum. i and  scutellum.  e r i o r notal-wing process i s prominent.  The f i r s t  s c l e r i t e i s p r o p o r t i o n a t e l y much longer appearing posteriorly.  The postaxillary to expand  The jugum, i f p r e s e n t , appears as a rounded  p o r t i o n of the wing a r t i c u l a t i n g with the t h i r d  axillary  s c l e r i t e which i s curved i n the opposite d i r e c t i o n . tegula i s larger.  The  The median suture i s a l i g h t - c o l o u r e d  line.  The metanotum ( P l a t e X, f i g . 3) i s s l i g h t l y s h o r t e r than the mesonotum. the t e g u l a are l a r g e r . mesonotum.  The t h i r d a x i l l a r y s c l e r i t e  The median suture i s as i n the  The jugum i s d e f i n i t e l y  absent.  and  -45Brachypterous R e p r o d u c t i v e s . * -  Zooterrnopsis angus t i collis„ The pronotum of a specimen,  which was d e r i v e d  d i r e c t l y "by the moulting of a seventh i n s t a r nymph, was smaller than that of a l a t e s .  The pronotum of a specimen  with  s h o r t e r wing pads ( P l a t e XIV, f i g . 9) was much longer and s l i g h t l y "broader.  The pronotum of a t h i r d specimen  i n which  the wing pads were c o n s i d e r a b l y longer and "broader than i n the other two  specimens was  A l l of these pronota may  longer "but of the same width.  be compared i n P l a t e V I I I , f i g . 3 ( 5 ) ,  P l a t e IX, f i g . 1 ( l and 2) and t a b l e 3. margin  In shape the a n t e r i o r  i s emarginate m e d i a l l y and the a n t e r o - l a t e r a l  are more acute than i n a l a t e s .  The pleuranotum  a n t e r i o r v e r t i c a l p o r t i o n are p r e s e n t . light-coloured.  margins  and the  The median suture i s  ITo dark spots are evident.  The mesonotum i s s i m i l a r to the drawing P l a t e  XV,  f i g . 1, of an apterous r e p r o d u c t i v e w i t h the exception that the rounded  sides are expanded i n t o wing pads.  suture i s a l i g h t - c o l o u r e d  The median  line.  The metanotum i s s h o r t e r and broader than the mesonotum. Apterous Reproductives• Zooterrnopsis a n g u s t i c o 1 1 i s . The nota of t h i 3 caste are quite simple ( P l a t e f i g s . 1 and 2 ) .  The v a r i a t i o n i n p r o n o t a l dimensions i s  XV,  -46i n d i c a t e d i n t a b l e 4 and the shape of the pronotum i s shown i n P l a t e VIII, f i g . 3(5).  I t s a n t e r o - l a t e r a l angles are  s t i l l more acute than i n the brachypterous forms. notum i s l o n g e r .  The l a t e r a l margins  metanotum are rounded. than the mesonotum.  The p r o -  of the mesonotum and  The metanotum i s broader and shorter  In other r e s p e c t s the nota are l i k e those  of the p r e c e d i n g c a s t e . Soldiers. Zootermopsis  angusticoliis.  The pronotum i s much the same i n shape as i n the a l a t e s but i s c o n s i d e r a b l y much l a r g e r ( P l a t e V I I I , f i g . 3 ( 2 ) . A brachypterous s o l d i e r had i t s pronotum intermediate i n shape between that of the normal tive.  The a n t e r i o r v e r t i c a l portion and the pleuranotum  those of a l a t e s . line.  s o l d i e r and apterous reproducare l i k e  The median suture i s i n d i c a t e d by a p a l e  The dark spots c h a r a c t e r i s t i c of a l a t e pronota are  absent.  In one specimen  the l e f t a n t e r i o - l a t e r a l angle was  produced  i n t o a l o b e , resembling s l i g h t l y a v e s t i g i a l wing. The mesonotum i s s l i g h t l y l e s s broad and i s longer  than the apterous mesonotum.  The l a t e r a l margins are t h i c k -  ened from the p o i n t where the p l e u r a l apodemes meet the notum to the a n t e r i o r margin.  In some cases the mesonotum may bear  v e s t i g i a l wings ( P l a t e XV, f i g s . 4 and 5 ) . The metanotum i s s h o r t e r than the mesonotum.  -47-  Zooterrnopsis  nevadensis.  '> The pronotum i s d i s t i n c t l y more angulate  anteriorly  i s a l s o much l a r g e r ( P l a t e V I I I , f i g . 4 ( 1 ) .  than i n the a l a t e and  In t a b l e 5 are given measurements of t h i s and the p r e c e d i n g species.  The a n t e r i o r margin of the pronotum i s s l i g h t l y  emarginate than i n a n g u s t i c o l l i a , difficulty  less  There i a a considerable  i n o b t a i n i n g uniform drawings or photographs of  the a n t e r i o r margin as witnessed by the work of others - due probably  to the f a c t that they are not viewed i n e x a c t l y  the same way.  In other d e t a i l s i t i s l i k e that of the pre-  ceding s p e c i e s . The mesonotum and metanotum are s i m i l a r to those in  angusticoilis. Reticuliterraes  hesperus.  In P l a t e IX, f i g . 2(3) the a n t e r i o r margin of the pronotum i s seen to be produced i n t o two The pronotum i s longer and broader,  lobes l a t e r a l l y .  t a p e r i n g much p o s t e r i o r l y .  The pleuranotum and the a n t e r i o r v e r t i c a l p a r t of the noturn are d i f f e r e n t i a t e d . a light-coloured  The median d o r s a l suture i s present  as  line.  The mesonotum and metanotum are each s h o r t e r than the pronotum, the mesonotum being longer than the metanotum. The l a t e r a l margins are not  thickened.  -48Immature Stages. * ' >  Zootermopsis  angusticoliis.  In newly hatched nymphs the pronotum and mesonotum are n e a r l y of the same s i z e , while the metanotum i s much s h o r t e r and s l i g h t l y broader.  The a n t e r i o r margin i s  not turned v e n t r a l l y and the pleuranotum able.  i s not d i s t i n g u i s h -  Median sutures a r e not y e t apparent. In nymphs of the t h i r d i n s t a r the three nota are  p r o g r e s s i v e l y broader p a s s i n g p o s t e r i o r l y ,  The pronotum and  mesonotum a r e approximately of equal l e n g t h and each l o n g e r than the mesonotum. distinct.  The pleuranotum  of the pronotum i s  Ho sutures on the meson a r e apparent.  THE PLEURA. Alates. Zootermopsis  angusticoliis.  The p l e u r a of the prothorax are of the g e n e r a l i z e d type.  The episternum extends from the noturn v e n t r a l l y almost  to the median l i n e  ( P l a t e V I I I , f i g s . 1 and 2 and P l a t e XI,  f i g . 4 ) , i s narrow a t both ends and i s broadest a t a l i t t l e above i t s middle p o i n t .  The epimeron  i s a l a r g e r , more or  l e s s rhomboid-shaped, s c l e r i t e extending down a l o n g s i d e the p o s t e r i o r margin of the coxal c a v i t y .  The trochant i n i s a  l o n g narrow s c l e r i t e a r t i c u l a t i n g with the a n t e r i o r margin of the coxa.  D o r s a l l y i t i s i n t i m a t e l y connected with a more  or l e s s small r e c t a n g u l a r s c l e r i t e .  This s c l e r i t e l i e s a l o n g  -49-  the p l e u r a l suture and with t h e e p i s t e r n u m . ;  p a r t of the tin.  On  seems to a r t i c u l a t e a n t e r o - d o r s a l l y F u l l e r (Sumner '33)  episternum.  claims  It seems to he a part  the d o r s a l margin of the coxa are two  extending a l o n g both s i d e s and apodeme meets the coxa. c o x i t e and  The  i t to "be a  of the  trochan-  narrow s c l e r i t e s  separated where the p l e u r a l  a n t e r i o r s c l e r i t e i s the b a s i -  the other the meron (Yfeber  '33).  The  coxella i s  s i t u a t e d p o s t e r i o r to the connection of the t r o c h a n t i n with the smaller  dorsal s c l e r i t e .  The  r e s t of the p l e u r a  e n t i r e l y of a membrane i n which the  ,5  prothoracic  n  consists  spiracle  i s s i t u a t e d near the caudal margin. Near the d o r s a l p a r t of the mesopleuron  and  metapleuron are s i t u a t e d the a l i f e r a e ( P l a t e X, f i g . 5 ) . medalifera  i s rectangular  i n shape and  elongate and rounded at i t s ends. epiBternum and The  The  the p o s t a l i f e r a i s d o r s a l t i p s of  the  epimeron are the p r e a l i f e r a and p l e u r a l i f e r a .  episternum i s l a r g e r than the epimeron and  i n i t s d o r s a l h a l f by a deep c l e f t . a rounded t i p , the coxal ular s c l e r i t e .  The  The  condyle.  trochantin  i s divided  V e n t r a l l y i t tapers The  epimeron i s a  c o n s i s t s of two p a r t s  to  rectangsepar-  ated by, an oblique l i n e so that the a n t e r i o r p o r t i o n i s crescent-shaped, the other t r i a n g u l a r .  The  coxella i s s i t -  uated i n the membrane near the p o s t e r i o r d o r s a l margin of the t r o c h a n t i n .  The meron (merum of F u l l e r ) i s a l a r g e  s c l e r i t e of the coxa proper.  It makes up  the d o r s a l p o r t i o n  -50af the coxa. prothoracic  The b a s i c o x i t e i s s l i g h t l y l a r g e r than i n the coxae.  The metapleuron i s e s s e n t i a l l y the same as the niesopleuron. Zootermopsis nevadensis. The a l a t e p l e u r a of t h i s species a r e s i m i l a r to those of a n g u s t i c o l i i s . Reticulitermes  hesperus.  The episternuia of the prothorax i s s l i g h t l y er than i n Zootermopsis. v e n t r a l l y ) than "broad.  The epimeron i s longer  species.  (dorso-  The t r o c h a n t i n and the small  d o r s a l to i t a r e of the same shape as i n the two  narrow-  sclerite  preceding  The "basicoxite and meron are s c a r c e l y v i s i b l e ,  although the i n t e r n a l r i d g e s e p a r a t i n g a n g u s t i c o l i i s i s present  them from the coxa i n  i n t h i s species.  The " p r o t h o r a c i c "  s p i r a c l e i s s u b c i r c u l a r i n o u t l i n e and i s r e l a t i v e l y The e p i s t e r n a of the mesothorax are s l i g h t l y longer,  smaller.  and metathorax  the d o r s a l p a r t of the epimeron i s  r e l a t i v e l y "broader, the meron i s l a r g e r and the t r o c h a n t i n c o n s i s t s of a s i n g l e s c l e r i t e . thoracic pleura.  The c o x e l l a i s absent i n a l l  The m e d a l i f e r a  i s a t r i a n g u l a r s c l e r i t e and  the p o s t a l i f e r a i s a long s c l e r i t e rounded a n t e r i o r l y and narrowed  to a point p o s t e r i o r l y .  The s t e r n o i d e a  i s a small  s c l e r i t e l o c a t e d more p o s t e r i o r l y than i n Zootermopsis.  -51Brachypterous  Reproductives.  Zooterrnopsis  ;  angusticoilis.  The p l e u r a of the prothorax alates. modified*  i s l i k e that i n the  The p l e u r a of the other two segments are somewhat The episternum i s completely  a n t e r i o r margin to the p l e u r a l suture.  s c l e r o t i z e d from i t s The epimeron i s very  prominent when viewed d o r s a l l y and v;ith the d o r s a l t i p of the episternum i t tapers to a p o i n t a t the d o r s a l p o i n t of the p l e u r a l apodeme which a r t i c u l a t e s w i t h a t h i c k e n i n g on the l a t e r a l margin o f the notum. p o s t a l i f e r a i s present ite.  In some specimens the  as a small p o o r l y d e f i n e d oval  scler-  The s t e r n o i d e a , both p a r t s of the t r o c h a n t i n and c o x e l l a  are s i m i l a r to those i n a l a t e s .  The meron i s a l s o a l a r g e  s c l e r i t e as i n the a l a t e s . Apterous  Reproductives.  Zooterrnopsis  angusticoilis.  The p l e u r a of t h i s caste i s l i k e those of the preceding  caste. Soldiers. Zo o t ermop s i s an gu s t i c o l l i s . The p l e u r a of a l l t h o r a c i c segments are l i k e  those  of apterous and brachypterous r e p r o d u c t i v e s except that the various s c l e r i t e s a r e s l i g h t l y l a r g e r and more h e a v i l y sclerotized.  =52Reticulitermes ;  hesperus.  The p l e u r a of t h i s caste a r e only s l i g h t l y more  s c l e r o t i z e d than those of workers and young repr-crductives. The p l e u r o n of the prothorax i s l i k e that of a l a t e s .  The  mesopleuron and raetapleuron a r e m o d i f i e d as i n Z, an gust i c o l l i s . The episternum  i s t r i a n g u l a r and completely  epimeron i s much the same as i n the a l a t e s .  sclerotized.  The  These two s c l e r -  i t e s taper d o r s a l l y , meeting at the d o r s a l t i p of the p l e u r a l apodeme, and a r t i c u l a t e w i t h a t h i c k e n i n g of the l a t e r a l margins of the nota.  The other p l e u r a l s c l e r i t e s a r e l i k e  those of a l a t e s . Immature. Zootermopsis  angusticoliis*  The p l e u r a of newly hatched  t e r m i t e s i s much  reduced d o r s o - v e n t r a l l y . The episternum and epimeron are e a s i l y d i s t i n g u i s h a b l e and a r e of the general shape of apterous r e p r o d u c t i v e s .  The meron of the meso- and meta-  thorax i s not y e t d i f f e r e n t i a t e d from the r e s t of the coxa. The trochant i n i s present as a s i n g l e crescent-shaped  sclerite.  STERNA Alates Zootermopsis The  angusticoliis.  sternum of the prothorax i s the shape of an  e q u i l a t e r a l t r i a n g l e ( P l a t e V I I I , f i g . 2, P l a t e XI, f i g . l ) .  -53It extends n e a r l y to the caudal margins of the coxal The mesosternum i s roughly  rectangular with i t s posterior  margin drawn to a rounded t i p . presternum*  cavities.  This i s much l a r g e r than the  I t has a median r i d g e on i t s e n t a l surface and  extends s l i g h t l y p o s t e r i o r to the a n t e r i o r margin of the coxal c a v i t i e s .  The sternum of the metathorax i s t r i a n g u l a r  w i t h i t s a n t e r i o r median p o r t i o n l a c k i n g . i n c a r e f u l l y stained preparations two  The metasternum  i s composed of three p a r t s ,  l a t e r a l and one small median p a r t  separated  former by a narrow s t r i p of membrane.  from the  This membrane  allows  f o r the s l i g h t f o l d i n g of the two l a t e r a l p a r t s over the median p a r t . of the coxal  The metasternum extends to the a n t e r i o r margins cavities. Zooterrnopsis  As  i n the preceding  nevadensis. species,  Reticulitermes The two  heBperus.  sternum of the prothorax i s represented  small elongated s c l e r i t e s which a r e separated  narrow p i e c e of membrane. the c o x a l  by  by a  These extend p o s t e r i o r l y between  c a v i t i e s ( P l a t e XI, f i g . 2 ) . The mesosternum i s t r i a n g u l a r w i t h the a n t e r i o r  median p o r t i o n membranous, thus roughly to the a n t e r i o r margins of the coxal  Y-shaped.  I t extends  cavities.  The metasternum i s much the same as that of the mesothorax but i s compressed a n t e r o - p o s t e r i o r l y .  -54Brachypterous and Apterous Reproductives. * •  Zootermopsis a n g u s t i c o l i i s . The presternum i s the same as i n the a l a t e s .  mesosternum i s e s s e n t i a l l y the same a n t e r i o r l y . it  The  Posteriorly  t a p e r s as i n the a l a t e s "but the l a t e r a l sides of the t i p  are equipped with l a r g e r setae than on the r e s t of the sternum. furca,  Posterior  to the sternellum, but a n t e r i o r  i s an oval p a t c h of l a r g e r s e t a e .  c o n s i s t s o f three p a r t s ,  to the  The metasternum  two l a r g e l a t e r a l wings separated  by membrane and one median p o s t e r i o r p o r t i o n to the t i p of the a l a t e metasternum.  corresponding  This median p o r t i o n ,  the basisternum l i k e that of the segment a n t e r i o r equipped l a t e r a l l y w i t h spines.  Between the  to i t , i s  invaginations  of the s t e r n a l apophyses i s another group of setae on the eusternum.  The p o s t e r i o r  t i p s of the sterna  of the meso-  and  metathorax are f a r more d i f f e r e n t i a t e d from the r e s t of  the  sternum than i n the a l a t e s . Soldiers. Zootermopsis a n g u s t i c o l l i s . The  sterna of t h i s caste a r e i d e n t i c a l w i t h those  of the two preceding  castes.  Reticulitermes The of the a l a t e s .  sterna  hesperus.  of the t h o r a c i c  segments are l i k e those  -55Immature '  Zooterrnopsis a n g u s t i c o i l i s . The  s t e r n a of newly hatched  t e r m i t e s a r e present  as p o o r l y d e f i n e d s c l e r i t e s and l i m i t e d to the v e n t r a l region proper, not extending l a t e r a l l y *  The presternum i s the l e a s t  s c l e r o t i z e d of a l l three sterna, i t s setae being l i m i t e d to two patches borders.  of small setae s i t u a t e d on the p o s t e r o - l a t e r a l  The s c l e r i t e i s v e n t r a l l y p l a c e d between the two  coxal c a v i t i e s .  The mesosternum i s b r o a d l y oval with many  l a r g e setae and extends s l i g h t l y between the coxal c a v i t i e s . The metasternum i s s h o r t e r , but of the same breadth, and equipped coxal  w i t h l a r g e setae, and extends l e s s between the  cavities.  SNDOSKSLETON. Alates. Zooterrnopsis a n g u s t i c o i l i s . E n d o s k e l e t a l s t r u c t u r e s of the nota, other phragmata a r e riot o f t e n found  i n insects.  than  In the pronotum,  extending between the a n t e r i o r d o r s a l p o r t i o n and the i n c u r v i n g a n t e r i o r more-ventral  p o r t i o n , there i s developed two  p i l l a r s of c h i t i n s i t u a t e d c l o s e r to the pleuranotum than to the meson ( P l a t e X I , f i g . 4 ) . V e n t r a l l y these p i l l a r s extend  l a t e r a l l y as t h i c k e n i n g s of the margin of the pronotum.  D o r s a l l y they a r e apparent  as dark spots and as s l i g h t de-  p r e s s i o n s of the notum proper, d e s c r i b e d above under nota. E n t a l l y the median suture i s present  as a s l i g h t r i d g e .  There  -56i s a l s o a t h i c k e n i n g a l o n g the meson a t the margin o f the incurved•portion.  In the mesonotum and metanotum there i s a  "broad v e r t i c a l p l a t e extending  t h r e e - f o u r t h s the d i s t a n c e  along the meson from the a n t e r i o r margins of the nota (Plate XI, f i g *  3)*  These p l a t e s are "broadest near the ant-  e r i o r margins and taper to a p o i n t p o s t e r i o r l y .  They give  to each of these nota c o n s i d e r a b l e r i g i d i t y and prevent "bending along a.ny l i n e p e r p e n d i c u l a r to .the meson. Phragmata occur on the a n t e r i o r margins of "both mesonotum and metanotum ( P l a t e X I I , f i g . 1 and P l a t e XI, f i g . 3).  The prephragma of the mesonotum i s quite deep and  i s s t r o n g l y emarginate.  L a t e r a l l y there are two w i n g - l i k e  p r o j e c t i o n s termed the y o l k of the prephragma (Sumner '33). The phragma of the metanotum i s only s l i g h t l y emarginate and does not extend  v e n t r a l l y to the same degree.  The y o l k i s  more prominent. The p l e u r a l endoskeletal s t r u c t u r e s c o n s i s t of l o n g curved apodemes ( P l a t e XI, f i g s . 4 and 5) and t h i c k e n ings.  The p r o p l e u r a l apodeme i s l i k e an i n v e r t e d Y with  the stem much elongated.  D o r s a l l y i t "bears on the v e n t r a l  s i d e of the p i l l a r w i t h i t s l a t e r a l extensions.  The apodeme  extends i n t e r n a l l y along the p l e u r a l suture to the d o r s a l margin of the coxal c a v i t y , becoming much broader r e a c h i n g the coxal condyle.  I t extends c e n t r a l l y to meet  the f u r c a e a r i s i n g from the prosternum. of the episternum  before  The a n t e r i o r margin  i s thickened i n t e r n a l l y .  The v e n t r a l t i p  -57of the t r o c h a n t i n has a long apodeme extending sally.  antero-dor-  The c o x e l l a a l s o has an apodeme p a s s i n g a n t e r i o r l y .  Beneath the suture s e p a r a t i n g the b a s i c o x i t e and the meron from the ooxa proper i s another r i d g e .  The p l e u r a l apodemes  of the mesothorax and metathorax are much longer and t h e i r extensions  r e l a t i v e l y not so prominent ( P l a t e X I , f i g . 5 ) .  There i s no t h i c k e n i n g of the a n t e r i o r margin of the e p i sternum.  There i s an i n t e r n a l r i d g e i n d i c a t e d e x t e r n a l l y by  the suture e x i s t i n g between the two p a r t s of the t r o c h a n t i n . In a d d i t i o n to the r i d g e of the d o r s a l p a r t of the coxa there i s a l s o a r i d g e extending an extension  from the coxal condyle as  of the l a t e r a l apodeme.  At the v e n t r a l t i p of  the t r o c h a n t i n there i s a small apodeme extending a n t e r i o r l y . The  c o x e l l a l apodeme i s present  i n these two t h o r a c i c  segments as w e l l . As i n other  i n s e c t s the endoskeleton of the  t h o r a c i c segments c o n s i s t s of the s t e r n a l apophyses or f u r c a l arms, the s t e r n e l l u m or furcasternum and the spina. the prothorax the s t e r n e l l u m i s absent.  The s t e r n a l apophyses  are r e l a t i v e l y s o f t l o n g i t u d i n a l p l a t e s extending coxal c a v i t i e s .  In  across the  The apodeme of the spinasternum, the spina,  i s w e l l developed and extends a n t e r i o r l y ( P l a t e XI, f i g . l ) . The mesosternal apophyses are much more h e a v i l y  scleritized  and about the same s i z e but do not extend to the p l e u r a l apodemes.  The spina i s s l i g h t l y l e s s elongate.  three processes  Between these  there i s a t h i c k e n i n g , the a n t e r i o r margin  -58of which gives r i s e to the sternellura.  This i s an  elongate  t h i c k e n i n g extending a n t e r i o r l y along the meson of the mesothorax except that the spina i s lacking,. Zooterrnopsis nevadensis The phragmata, apodemes and apophyses of each t h o r a c i c segment are the same.  The  unusual t h i c k e n i n g s of  the nota d e s c r i b e d i n a n g u s t i c o i l i s are present  in this  species as w e l l . R e t i c u l i t e r m e s hesperus„ The n o t a l e x s k e l e t a l s t r u c t u r e s other than the phragmata are much reduced i n t h i s s p e c i e s .  The  two  thicken-  ings o c c u r r i n g between the a n t e r i o r p a r t s of the pronotum are reduced to t h i c k e n i n g s of the v e n t r a l margin of the a n t e r i o r v e r t i c a l p o r t i o n adjacent the p r o p l e u r a l apodemes.  to the d o r s a l ends of  There i s a s l i g h t t h i c k e n i n g of  the a n t e r i o r t i p of the median suture on the v e r t i c a l face of the pronotum.  The phragmata of the mesothorax and meta-  thorax are concave a n t e r i o r l y .  The y o l k i s not  distinct.  In the p l e u r a the l a t e r a l apodemes are more elongate,  the inner p r o j e c t i o n of i t being d i s t i n c t l y more  slender than that of Z. a n g u s t i c o i l i s .  The apodemes of the  other t h o r a c i c segments are l i k e w i s e slender, the inner p r o j e c t i o n b e i n g l o n g and  tapering.  The apodemes from the  inner surface of the t r o c h a n t i n s of the p o s t e r i o r two t h o r a c i c p l e u r a are not present.  The a n t e r o - d o r s a l margin of  -59each coxa has an i n t e r n a l r i d g e present p a r t i c u l a r l y i n the mesothorax and metathorax.  The t h i c k e n i n g along the coxo-  meron suture appears as an extension of the p l e u r a l apodeme. V e n t r a l l y the endoskeleton of the prothorax i s poorly defined.  The f u r c a l arms a r e t h r e a d - l i k e and l o n g .  The s p i n a i s l i t t l e i n Zootermopsis  s c l e r o t i z e d and extends a n t e r i o r l y as  ( P l a t e XI, f i g . 2 ) .  In the mesothorax, the  f u r c a l arms a r e more slender than i n the two.preceding species the s p i n a "being r e l a t i v e l y l a r g e r and the s t e r n e l l u m "being p a r t i c u l a r l y prominent.  The endoskeleton of the metasternum  i s i d e n t i c a l w i t h that of the p r e c e d i n g segment except  that  the spina i s absent* Brachypterous and Apterous Reproductives. Zootermopsis  angusticoliis.  The endoskeleton of these castes i s not so apparent as i n the a l a t e s because f i g . 3).  of l e s s s c l e r o t i z a t i o n ( P l a t e XII  The p i l l a r s c h a r a c t e r i s t i c o f a l a t e pronota a r e  reduced to t h i c k e n i n g s adjacent to the p r o p l e u r a l apodeme3. The mesonotum and metanotum both l a c k the median p l a t e s . There are t h i c k e n i n g s along the a n t e r o - l a t e r a l margins mesonotum and metanotum.  of the  The phragmata are much reduced  ( P l a t e XV, f i g . 1 and 2 ) .  The p r o p l e u r a l apodemes are l i k e  those of a l a t e s but the l a t e r a l apodemes of the other segments a r e intermediate between the p r o t h o r a c i c apodemes and the mesothoracic apodemes of a l a t e s ; that i s , they a r e not as l o n g and the inner p r o j e c t i o n i s broader.  Apodemes occur  -60on the t r o c h a n t i n s of the meso- and metathorax on the e p i sternum of the prothorax and on the c o x e l l a .  Thickenings or  r i d g e s are present on the episternum of the prothorax, the a n t e r o - d o r s a l margins of the coxa a l o n g the suture separa t i n g the coxa proper from the meron, and along the suture s e p a r a t i n g the p a r t s of the t r o c h a n t i n . i s s l i g h t l y modified.  The s t e r n a l skeleton  The spine of the prosternum  i s just  v i s i b l e and that of the mesosternum i s i d e n t i c a l with that of a l a t e s .  The s t e r n a l apophyses are much the same and  s t e r n e l l a are s c a r c e l y  the  visible.  Soldiers. Zooterrnopsis a n g u s t i c o i l i s and nevadensis„ The t h o r a c i c endoskeleton of t h i s caste i s e s s e n t i a l l y the same as that of the two p r e v i o u s c a s t e s .  The  a n t e r o - l a t e r a l t h i c k e n i n g s of the mesonotum and metanotum are more prominent  and the f u r e a l arms are s l i g h t l y  Reticulitermes  larger.  hesperus.  With the exception of the mesosternal  apophysis,  the endoskeleton i s l i k e that of a l a t e s but i s not so h e a v i l y sclerotized. centrally  I t has a dark d o r s a l p o r t i o n a r i s i n g from i t  (the spina) with the caudal p o r t i o n of the whole  s t r u c t u r e t e r m i n a t i n g i n a rounded p r o j e c t i o n .  -61Immature. •  Zooterrnopsis a n g u s t i c o i l i s . In newly hatched  nymphs the p l e u r a l apodemes are  by the f a r the most developed endoskeleton. extremely  The  faint.  i n the prosternum. as widely  s t r u c t u r e s of the t h o r a c i c  s t e r n a l e n d o s k e l e t a l s t r u c t u r e s are Two  small f u r c a l arms can be d i s t i n g u i s h e d  The.mesothoracic f u r c a l arms are v i s i b l e  separated i n v a g i n a t i o n s .  those of the previous segment. sternum are even more widely  These are much l a r g e r than  The f u r c a l arms of the meta-  separated.  Other e n d o s k e l e t a l  s t r u c t u r e s are s c a r c e l y i f at a l l developed* LEGS. Alates. Zooterrnopsis a n g u s t i c o l l i s The  three p a i r s of l e g s are almost  identical,  d i f f e r i n g c h i e f l y i n s i z e and p r o p o r t i o n and i n the absence or presence of the meron.(Plate X, f i g , 5 and P l a t e XII, fig.  5). The  coxae of the prothorax  b e i n g longer than broad.  On  are  subcylindrical,  the a n t e r o - d o r s a l margins are  s i t u a t e d the narrow b a s i c o x i t e and meron ( P l a t e V I I I , f i g s . 1 and 2 ) .  The other coxae are much shorter and broader  and are  d i s t i n g u i s h e d by the l a r g e meron on the d o r s a l s u r f a c e . t r o c h a n t e r i s small and between the coxa and  subtriangular.  The  In the membrane  trochanter there are three small access-  ory s c l e r i t e s  (Stunner *33).  as long as the coxa and "broad. The  The femur i s approximately  i s n e a r l y four times as long  as  The mesofemur i s s h o r t e r than e i t h e r of the other  t i b i a i s shortest i n the p r o l e g s and  ,legs.  twice  Each t i b i a bears a f a i r l y  longest  constant  two  i n the meta-  number of  spines.  The p r o t h o r a c i c l e g s have seven spines, of which f o u r are a p i c a l , two and  one  are on the d o r s a l s u r f a c e near the middle part  on the v e n t r a l p a r t near the d i s t a l end.  t h o r a c i c l e g has (one  three a p i c a l spines on the v e n t r a l surface  of these i s s t r i c t l y  subapical) and  s u r f a c e near the middle of the t i b i a . has  s i x , of which two  are a p i c a l and  ated on the v e n t r a l s u r f a c e .  two  on the d o r s a l  The metathoracic  are c h a r c t e r i s t i c a l l y barbed or saw margins ( P l a t e X I I I , f i g . 4 ) .  The  j o i n t e d , the second most proximal  d r i c a l , the t h i r d and  toothed  spines lateral  being wedge-shaped and  only  The b a s a l t a r s a l segment i s c y l i n  f o u r t h l i k e s e c t i o n s of a c y l i n d e r cut  f o u r combined.  The  than the  other  f i f t h segment or p r e t a r s u s i s narrower  segments p r o x i m a l l y but  i s as broad  l a r g e claws with a median pad  (Plate XIII, f i g . 5). The  The  the  t a r s i are always f i v e -  the f i f t h i s s l i g h t l y longer  the u n g u i f e r .  Usually  on t h e i r  o b l i q u e l y while  It bears two  separ-  There i s often v a r i a t i o n from  number o f spines v a r i e s between f i v e and n i n e .  slightly visible dorsally.  leg  the remainder are  these d e s c r i p t i o n s even on the same i n d i v i d u a l .  than the other  The meso-  distally.  or p u l v i l l u s  Each claw i s a r t i c u l a t e d d o r s a l l y to  arolium,  l i k e the claws, i s hollow and i s  -63a direct "base.  c o n t i n u a t i o n of the median p a r t of the p r e t a r s a l  I t ' i s e n t i r e l y membranous.  The u n g u i t r a c t o r  i s a large  median p l a t e which i s p a r t l y i n v a g i n a t e d  i n t o the end  of  tarsus.  the l o n g and  tapering  To i t s proximal  end i s attached  tendon of the r e t r a c t o r muscle of the  the  claws.  Zootermopsis nevadensis. The l e g s of t h i s species are l i k e those of a n g u s t i c o l i i s except that v a r i o u s h a i r s on the upper of the hind t i b i a are f u l l y one-half  surface  the diameter of the tibia„  In a n g u s t i c o l l i s they are l e s s than h a l f . R e t i c u l i t e r m e s hesperus„ As i n Z. a n g u s t i c o l i i s a l l l e g s are much a l i k e (Plate XII, f i g . 6). but  The meron i s i n d i s t i n c t  i s l a r g e i n the other  two.  i n the p r o l e g  The mesothoracic and meta-  t h o r a c i c coxae are much l a r g e r than those of the The  prothorax.  femur of the mesoleg i s smaller than the other two  the t i b i a of the p r o l e g i s smaller than the o t h e r s . l e g has three a p i c a l spines on the t i b i a , v e n t r a l and the other  the other d o r s a l .  two  There are two  The  pro-  of these are a p i c a l spines  two p a i r s of l e g s , both being v e n t r a l .  are s l i g h t l y toothed.  and  These  on  spines  There are only four t a r s a l segments  with the second, the s m a l l e s t , and wedge-shaped.  The  first  three have protuberances developed d i s t a l l y on the v e n t r a l side. first  These are l a r g e s t on the t h i r d and t a r s a l segment.  smallest on  the  A l l p a r t s of the l e g s are covered  with  -64numerous small setae.  In l e n g t h , those of the tarsus are  equal to or greater than the -width of the t a r s u s ; those of the t i b i a ,  one;-.half the width of the t i b i a ;  femora o n e - t h i r d to one-quarter  those of the  the width of i t .  The  pre-  t a r s u s d i f f e r s i n some r e s p e c t s from that found i n Zooterrnopsis.  The u n g u i t r a c t o r p l a t e i s r e l a t i v e l y shorter,  the a r o l i u m i s present as a knob, one the u n g u i t r a c t o r p l a t e and has  t h i r d the l e n g t h of  s i t u a t e d at i t s base a median  seta.  Brachypterous  Reproductives.  Zooterrnopsis angusticollis„ Of the three p a i r s of l e g s , the metaleg i s the l a r g e s t , as i n the a l a t e s . than those of the a l a t e s .  They are s l i g h t l y s t o u t e r and  longer  The femora of the f i r s t p a i r are  twice as wide as the t i b i a , of the second p a i r two and  one-  h a l f times, and of the t h i r d p a i r three times.  spines  are present as i n the a l a t e s but with one  Tibial  exception of the  a p i c a l spines, they are smaller than those of the a l a t e s . tibial  spines of the p r o l e g are l o c a t e d as f o l l o w s : two  The  large  a p i c a l v e n t r a l , one l a r g e a p i c a l d o r s a l , one l a r g e a p i c a l ( s l i g h t l y smaller than the others) inner d o r s o - l a t e r a l . spines are s e r r a t e d but  the number of s e r r a t i o n s i s l e s s  than i n the a l a t e s , there b e i n g f i v e or s i x . l a r g e a p i c a l v e n t r a l , one small at l / 2 and 7/8  Apical  Mesoleg:' two  s l i g h t l y l a r g e r a p i c a l d o r s a l , two  v e n t r a l , three small at 3/8,  l / 2 and  5/8  -65 = dorsal.  Me'taleg:  two l a r g e a p i c a l v e n t r a l , three small  ( l / 3 si'ze of the a p i c a l ) at l / 2 , 3/4 and 7/8 v e n t r a l . h a i r s are one-fourth  the width of the t i b i a .  femora are of the same l e n g t h .  Tibial  Those of the  As i n the a l a t e s there i s ,  at one-fourth the d i s t a n c e from the f e m o r a l - t i b i a l j o i n t on the d o r s a l s i d e of the t i b i a , a s l i g h t rounded p r o j e c t i o n which i s present  i n a l l three p a i r s of l e g s .  The t a r s a l  segments are s i m i l a r to those of the a l a t e s , the second, t h i r d and f o u r t h being membranous at the d i s t a l  end.  a r o l i u m or pad l o c a t e d between the claws i s reduced very s l i g h t rounded protuberance Apterous  and i s probably  The  to a  funtionless„  Reproductives.  Zootermopsis a n g u s t i c o l i i s . The l e g s of t h i s caste are the same s i z e as those of the p r e v i o u s l y d e s c r i b e d c a s t e . different  i n number although  t h i s may be a v a r i a t i o n .  a p i c a l inner dors'o-lateral, present p r o l e g i s absent. same.  The spines are somewhat The  i n the brachypterous  The other spines of the p r o l e g are the  Of the mesoleg there i s a small a p i c a l d o r s a l , two  small v e n t r a l at l / 2 and 5/8 and one small d o r s a l at 5/8. Of the metaleg there are small setae at l / 2 , 5/8 and 7/8, and a s u b a p i c a l v e n t r a l .  A l l the setae are s e r r a t e d .  Soldiers. Zootermopsis a n g u s t i c o l l i s . The l e g s are much l a r g e r than i n any of the other forms and i n the case of the t i b i a the shape of the proximal  -66end  i s different  ( P l a t e X I I I , f i g . l ) . The femora a r e  n o t i c e a b l y much broader.  On the v e n t r a l s i d e of the t i b i a of  the metaleg there a r e two l a r g e crenulated smaller  s u b a p i c a l 3 p i n e and two other  spines, one  spines of the same  l e n g t h aa the l a t t e r l o c a t e d at 1/2 and 5/8.  The mesoleg has  i n a d d i t i o n three d o r s a l spines l o c a t e d at l / 2 , 3/4 and apical.  The p r o l e g has the two p a i r e d l a r g e a p i c a l s and three  d o r s a l s as i n the mesoleg.  The p r e t a r s u s  d i f f e r s from that  of a l a t e s i n that the claws a r e much stouter, the u n g u i t r a c t o r p l a t e i s quite d i f f e r e n t  i n shape, being broader at i t s  middle and t a p e r i n g to a blunt p o i n t p r o x i m a l l y  and an  arolium which has almost disappeared although i t has a spine on i t s v e n t r a l surface  (Plate XIII, f i g . 6 ) .  Zooterrnopsis The species. has  nevadensis.  l e g s a r e much the same as i n the preceding  The t i b i a l  setae are l o c a t e d as f o l l o w s : the p r o l e g  three l a r g e a p i c a l ,  two of these v e n t r a l and the other  i s outer l a t e r a l , one smaller  s u b a p i c a l d o r s a l and one small  •at l / 2 and 5/8; the mesoleg has two l a r g e a p i c a l v e n t r a l , one  3 m a i l s u b a p i c a l v e n t r a l , one small subapical  dorsolat-  e r a l , one small v e n t r a l at l / 2 and three small l a t e r a l at l / 2 , 5/8 and 3/4 (or as i n the other mesoleg two small l a t e r a l at l / 2 and 3/4);  the metaleg has two l a r g e a p i c a l  v e n t r a l and three smaller v e n t r a l at l / 2 , 3/4 and  tf/8j  -67Reticulitermes  hesperus.  Of the three l e g s the metaleg i s the l a r g e s t c u l a r l y w i t h respect intermediate  to the l e n g t h of the t i b i a ) ,  (parti-  the p r o l e g i s  i n s i z e w i t h the femora as l a r g e as that of the met-  a l e g ( s l i g h t l y narrower) hut the t i b i a i s much shorter  (Plate  X I I I , f i g . 3 ) . The femora of the mesolegs are smaller  than  those of the other  two p a i r s o f l e g s .  The t i b i a of the mesoleg  i s of the same l e n g t h as that of the p r o l e g .  The legs  differ  from a l a t e l e g s i n "being not so slender.  A l l have two a p i c a l  ventral t i b i a l  tarsus.  spurs and a four-3egmented  The p r e t a r -  s i s i s e x a c t l y the same as that of a l a t e s except that the a r o l ium  i s g r e a t l y reduced and the claws are s l i g h t l y  serrated.  Immature. Zootermopsis a n g u s t i c o l i i s . The  l e g s of newly hatched nymphs a r e much d i f f e r e n t i n  form and t h e i r s e t a l v e s t i t u r e ( P l a t e X I I I , f i g . 2 ) . The femur i s two and one-half s l i g h t l y longer  times the l e n g t h of the trochanter,  the t i b i a  than the femur and the tarsus as long as the  femur.  The femur i s s l i g h t l y broader than the other  ments.  The l e g s a r e c l o t h e d with long slender  l e g seg-  setae, at l e a s t  as l o n g as the width of the t i b i a and in,some cases twice i t s width.  The two larger-ventral a p i c a l setae c h a r a c t e r i s t i c of the  t i b i a e of other and  castes a r e reduced to very small cone-shaped  the f o u r t h i 3 t u b e r c u l a t e  d i s t a l l y . . The i n g u i t r a c t e r p l a t e  i s t r i a n g u l a r , smaller and i s not developed d i s t a l l y as i n alates.  The arolium  i s much reduced with a l o n g median seta.  -68THB WINGS. * -Alates. Zootermopsis a n g u s t i c o l l i s and nevadensis. Under~ t h i s heading w i l l "be described  the a r t i c u l -  a t i o n s of the wings "by the wing s c l e r i t e s or p t e r a l i a e and the wing  venation. The p t e r a l i a e proper, that i s , the a x i l l a r y  have "been d e s c r i b e d above under the n o t a (page 43). s c l e r i t e s a r t i c u l a t e against and  against  sclerites, These  the n o t a l wing processes d o r s a l l y  the a l i f e r a e of the p l e u r a .  Bearing  on the d i s t a l  s u r f a c e s of the a x i l l a r y s c l e r i t e s are the proximal ends of the v e i n s , named from the v e i n s of which they are p r o j e c t i o n s . Of these,  three are d i s t i n c t : the c o s t a l i s , r a d i a l i s and  a n a l i s ( P l a t e X, f i g . l ) . The  two p a i r s of wings a r e nearly of equal s i z e ,  those of Z. nevadensis being  s l i g h t l y smaller  (Plate XIII,  f i g . 7 and Tables 1 and 2 ) . They are about twice the length of the body and about three times as l o n g as broad and a r e densely r e t i c u l a t e w i t h the margins not c i l i a t e . wings a r e broken o f f a f t e r f l i g h t . a c l e a r l y d e f i n e d humeral suture  The hind wings do hot have  so that the f r e e edge of the  hind wing stub i s sometimes i r r e g u l a r . are much longer  The f r o n t wing scales  than the pronotum and u s u a l l y twice as l a r g e  as the hind p a i r . f i g . 7.  The f o r e  The wing venation  The c o s t a l v e i n i s l a c k i n g .  i s shown by P l a t e XIII, Subcosta i s u s u a l l y  separate from r a d i u s and extends to the f i r s t  f o u r t h of the  -69wing.  Radius gives r i s e to three "branches: r a d i u s one,  two p l u s three and r a d i u s four p l u s f i v e . r a d i u s i n the wing s c a l e ( P l a t e XIV,  Radius one  radius  leaves  f i g . l ) , appearing  in a  few wings to coalesce with r a d i u s two p l u s three proximally or with subcosta d i s t a l l y * branched but  i n one  Radius two p l u s three u s u a l l y i s un-  case i t f o r k e d d i s t a l l y .  There are many  d i s t a l branches of r a d i u s four p l u s f i v e given o f f a n t e r i o r l y , a l l h e a v i l y s c l e r o t i z e d so that the support i s mostly  by t h i s v e i n and  i t s branches.  given by the veins  Media i s not  so  prominent and p a s s e s f through the c e n t r a l p a r t of the wing. has three or f o u r branches. media and  i s two  Cubitus coalesces p r o x i m a l l y with  or three-branched.  only on the wing s c a l e .  It  The anal v e i n i s present  Between the v e i n s media p l u s cubitus  and a n a l there i s a l i n e of weakness. In the h i n d wing f i g u r e d ( P l a t e XIV, one  i s q u i t e d i s t i n c t with subcosta u n i t i n g with i t p r o x i m a l l y .  The f i r s t  branch of the r a d i a l  cond coalesces p a r t way and  f i g . 2), radius  sector i s unbranched and the se-  with media.  the a n a l v e i n unbranched.  Cubitus i s six-branched  The hind wing l a c k s the humeral  suture. R e t i c u l i t e r m e s hesperus. The v e n e l l a e or proximal  ends of the veins are not  at a l l d i s t i n c t . As i n Zooterrnopsis both p a i r s of wings are almost identical of  i n s i z e and venation and are about twice the length  the body.  The humeral suture i s w e l l d e f i n e d i n both wings  -70of R e t i c u l i t e r i n e s whereas i n Zoot ermopsis the suture i s very indistinct  i n the h i n d wings.  The suture i n the forewings i s  more d i s t a l than i n the hind wings.  The wings show consider-  a b l e r e d u c t i o n of v e i n s , only r a d i u s and median being present as simple unbrarrhsd v e i n s . radius four plus f i v e . which those nearer  Radius according to Comstock i s  Cubitus bears  s e v e r a l branches of  the humeral suture are more prominent.  area occupied by cubitus and i t s branches i s n e a r l y the t o t a l area of the wing.  The  one-half  As i n Leucotermes the anal area  i s r e s t r i c t e d to the scapular s h i e l d .  There are no anal v e i n s .  Between the l a r g e r veins there i s a network of w r i n k l e s . Brachypterous  Reproductives.  Zootermopsis a n g u s t i c o l i i s . In t h i s caste, the l o s s of complete development of wings i s c o r r e l a t e d with the l o s s of the p t e r a l i a e . The wing pads c l o s e l y resemble those of the seventh i n s t a r nymphs ( P l a t e XIV, f i g s . 6, 8 and 9 ) . There i s much v a r i a t i o n i n the l e n g t h of these wing pads as shown by the i l l u s t r a t i o n s and Table 3.  The wing i l l u s t r a t e d i n f i g u r e 6  was of an i n d i v i d u a l which became s e x u a l l y mature a f t e r moult i n g from a t y p i c a l seventh  i n s t a r nymph.  In comparing i t with  the wing pads of such nymphs ( f i g . 7), the pad i s seen to be s l i g h t l y longer.  In the other f i g u r e s , the wing pads are of  i n d i v i d u a l s whose l i f e h i s t o r y was not known.  The wing pad  f i g u r e d i n 8 was f o l d e d across i t s l e n g t h and was much l a r g e r than the seventh  i n s t a r wing pad.  In s i m i l a r  individuals  -71as f i g u r e d i n 8 the pads were curved down along the sides of the thorax.  F i g u r e 9 i s the wing pad of an i n d i v i d u a l taken  from a l a r g e colony.  It i s quite small.  A l l of these wing  pads have tracheae s i m i l a r l y p l a c e d as i n seventh i n s t a r nymphs ( P l a t e XIV, f i g s . 3 and 4 ) . Apterous Reproductives. In t y p i c a l apterous r e p r o d u c t i v e s , the wing pads are represented by f o l d i n g s of the l a t e r a l edges of the mesonotum and metanotum ( P l a t e XV, f i g . l ) .  In others, t h i s f o l d i n g  becomes deepened so that i t appears as a v e s t i g i a l wing ( P l a t e XV, f i g . 2 ) .  There i s much v a r i a t i o n amongst i n d i v i d -  u a l s and even on the s i d e s of one specimen.  With a large;num-  ber of t h i s and p r e c e d i n g castes, probably a complete s e r i e s showing the wing-pad development could be found. Soldiers. Zootermopsis a n g u s t i c o i l i s . In t h i s caste the f o l d i n g of the l a t e r a l  sides of  the mesonotum and metanotum i s not so broad ( P l a t e XV, f i g . 3). In some i n d i v i d u a l s which may be s e x u a l l y mature there i s some wing-pad development ( P l a t e XV, f i g s . 4 and 5 ) .  As i n  apterous forms there i s much v a r i a t i o n between specimens and even on d i f f e r e n t  s i d e s of the same i n s e c t .  Wing pads of the  s i z e i l l u s t r a t e d i n f i g u r e 4 or s l i g h t l y smaller have the three p r i n c i p a l tracheae extending from the a n t e r o - l a t e r a l margin of the hoturn i n t o the wing pad.  These wing pads are  shorter than those o c c u r r i n g on seventh i n s t a r nymphs. specimen on the l e f t  In one  side of the mesonotum there was an un-  -72m u t i l a t e d p r o j e c t i o n of the v e n t r a l margin ( P l a t e XV, f i g . 5). R e t i c u l i t e r m e s hesperus. The  s o l d i e r s examined had no i n d i c a t i o n s of wing  pads. Immature. Zootermopsis a n g u s t i c o l l i s . In seventh i n s t a r nymphs the t r a c h e a t i o n of the wing pads c l o s e l y resembles the venation which the l a t t e r was d e r i v e d  of the a d u l t s from  ( P l a t e XIV, f i g . 3 ) .  Subcosta  i s two-branched with the a n t e r i o r "branch short and not shown i n the i l l u s t r a t i o n .  Radius gives o f f two f a i n t  tracheae,  r a d i u s one and r a d i u s two p l u s three, near the base of the wing pad.  The other branch r a d i u s four p l u s f i v e i s a l a r g e  fteachea w i t h many f i n e r tracheae  branching  as l a r g e and prominent as the preceding tracheae  branching  from i t p o s t e r i o r l y .  u s u a l l y six-branched.  from i t .  and has three  Media i s fine  The cubitus area i s  The anal trachea a r i s e s from the same  trachea as cubitus but i s l i m i t e d to the proximal part of the f o r e wing. THE ABSOMEH. Alates. Zootermopsis a n g u s t i c o l i i s . The 3ternites.  abdomen c o n s i s t s of abdominal t e r g i t e s and nine  The small s c l e r i t e shown i n P l a t e XI, f i g . 1,  -73posterior  to the  i n v a g i n a t i o n s which form the f u r c a l arms of  the metathorax i s the v e s t i g i a l f i r s t the f i r s t  t e r g i t e a l l the others are  posterior  two  or three are  l a c k s the a n t e r o - l a t e r a l four  sternite.  Apart from  s i m i l a r except that  s l i g h t l y smaller.  The  first  the tergite  corners, i t s s p i r a c l e i s three or  times as l a r g e as those of the other t e r g i t e s , and  a narrow phragma ( P l a t e XII,  f i g . l ) . A l l hut  i t has  the n i n t h  and  t e n t h s t e r n i t e s possess s p i r a c l e s . In the male s t e r n i t e s two ( P l a t e XVI,  f i g . 2).  Posterior  p a r a p r o c t s separated on the n i n t h  to nine are  to the n i n t h  the median l i n e .  On  i s a pair the  of  caudal end  of  s t e r n i t e are a p a i r of s t y l e t s or subanal s t y l e s .  A p a i r of f i v e - j o i n t e d c e r c i are l o c a t e d t e r g i t e ten and portion  simple p l a t e s  i n the membrane "between  the paraproct j u s t v e n t r a l  of t e r g i t e  to the  anterior  ten.  In the female, the  s t e r n i t e s from two  s i m i l a r to those of the male.  Sternite  to s i x  seven i s much elongated  c o v e r i n g the much m o d i f i e d eighth s t e r n i t e ( P l a t e XVI, Subanal s t y l e s are absent.  The  The  c e r c i are  the  eighth s t e r n i t e i s shown by  paraprocts are  s i m i l a r to those of the males.  Three small p a r t s are lying posterior  to the opening of the  p a r t s on e i t h e r  side of the  v e n t r a l v a l v u l a e are D o r s a l to the  ninth  supported by  v e n t r a l v a l v e s and  fig.l ) .  s l i g h t l y longer. The  f i g u r e s 3, 4 and  s c l e r o t i z e d , one  are  character of 5 of P l a t e  a median v e n t r a l oviduct and  sternite.  The  XVI.  portion  the other  two  lobe-like  the narrow bars, the v a l v i f e r s . situated  i n the n i n t h  sternite  -74i s a V-shaped s c l e r i t e r e p r e s e n t i n g the v e s t i g e s of the  two  slender' "bars or i n f e r i o r apophyses of the d o r s a l v a l v u l a e (Walker  '19).  gootermopsis The s t e r n i t e s angusticoliis.  nevadensis.  of "both sexes are l i k e those of  The g e n i t a l i a of the female are i d e n t i c a l .  Anal s t y l e t s are l a c k i n g i n the female. are  Those i n the male  r e l a t i v e l y l a r g e r than those of a n g u s t i c o l i i s . R e t i c u l i t ermes hesperus. The abdomen c o n s i s t s ^ o f ten t e r g i t e s and nine  sternites,  the f i r s t  sternite  The s p i r a c l e of the f i r s t abdominal segments.  i s v e s t i g i a l ( P l a t e XI, f i g . 2 ) .  segment i s l i k e those of the other  The c e r c i are two-jointed, the b a s a l  j o i n t b e i n g twice as broad as long, and the terminai twice as l o n g as broad and t a p e r i n g to -a b l u n t t i p . In peared.  the female, the g e n i t a l i a have e n t i r e l y d i s a p -  The subanal s t y l e t s are absent as i n Zootermopsis.  The seventh s t e r n i t e o v e r l a p s the e i g h t h and part of the n i n t h . The male g e n i t a l i a are not developed. present on the n i n t h s t e r n i t e . of  S t y l e s are \ , / ... .1  The s t e r n i t e s are l i k e those j  Zootermopsis. Brachypterous and Apterous Reproductives• Zootermopsis  angusticollis.  The abdomen i s much l i k e that of s e v e n t h - i n s t a r  /lij-,^ "' 1  0  nymphs except that i t i s s l i g h t l y l a r g e r and more h e a v i l y sclerotized.  Anal B t y l e t s a r e present i n "both sexes*  The  number of v i s i b l e s t e r n i t e s i n the female i s the same as i n the alates.  The v a l v u l a e and the h e a v i l y s c l e r o t i z e d p a r t s of the  g e n i t a l i a a r e s i m i l a r to those of the a l a t e s . of the male are l i k e those of male a l a t e s . segmented.  The s t e r n i t e s  The c e r c i are f i v e -  The abdomen has eight s p i r a c l e s ,  the f i r s t i s  twice as l a r g e as the others and i s hidden i n l i v e specimens by the epimeron  of the t h i r d t h o r a c i c  segment.  R e t i c u l l t ermes hesperus * In other s p e c i e s of the same genus the anal s t y l e t s are absent  i n the females (Thompson and Synder *20). Soldiers. Zooterrnopsis a n g u s t i c o i l i s and nevadensis. The abdomen of s t e r i l e s o l d i e r s i s smaller than that  of apterous r e p r o d u c t i v e s .  In others i t may be nearly equal  i n s i z e although u s u a l l y not so broad. men has five-segmented  The t i p of the abdo-  c e r c i and a p a i r of anal s t y l e t s .  When  female s o l d i e r s do become f e r t i l e there are no external chara c t e r s such as the v a l v u l a e which d i f f e r e n t i a t e them. R e t i c u l i t e r m e s hesperus. T h i s species has two-segmented c e r c i and anal f i l a ments.  -76Immature. *  Zooterrnopsis In newly hatched  angusticoilis.  nymphs, the subanal  r e l a t i v e l y much longer and a r e two-segmented. as long as the unsegmented c e r c i and extend of the l o b e - l i k e p a r a p r o c t s .  s t y l e t s are They are nearly  almost to the t i p s  These l a t t e r are widel3  f  separ-  ated and are c l o t h e d with l a r g e setae. In t h i r d - i n s t a r nymphs the b a s a l p a r t of the s t y l e s appears as a protuberance are r e l a t i v e l y s h o r t e r . meeting on the meson.  of the n i n t h s t e r n i t e .  The s t y l e s  The paraprocts are broad p l a t e s The c e r c i are s t i l l  unsegmented.  INTERNAL ANATOMY. A comparison of the Three Species. THE DIGESTIVE TRACT. . Zootermopsis a n g u s t i c o l l i s . As i n a l l i n s e c t s the alimentary embryonically  canal c o n s i s t s  of three w e l l - d e f i n e d r e g i o n s : the f o r e i n t e s -  t i n e or stomodaeum, the mid i n t e s t i n e or mesenteron and the hind i n t e s t i n e or proctodaeurru proctodaeum are formed embryonically  The stomodaeum and as i n v a g i n a t i o n s of the  a n t e r i o r and p o s t e r i o r body w a l l r e s p e c t i v e l y  and';therefore  are l i n e d with the same m a t e r i a l as that of the that i s c h i t i n .  (stomach)  exoskeleton,  The mid i n t e s t i n e , a r i s i n g independently  the body w a l l , l a c k s t h i s c h i t i n o u s l i n i n g .  of  The d i g e s t i v e  t r a c t Is the same f o r a l l c a s t e s , v a r y i n g .only i n s i z e and the amount of food which i t c o n t a i n s .  Newly-hat ched nymphs  u s u a l l y have a l a r g e bubble of a i r i n the abdominal p o r t i o n of the alimentary  canal.  In more mature i n d i v i d u a l s p a r t s of  the d i g e s t i v e t r a c t may be enlarged before  ecdysis,  containing  nothing but a i r . Stomodaeum. Food on e n t e r i n g the p r e o r a l c a v i t y ) p a s s e s through the mouth i n t o the buccal the d i g e s t i v e t r a c t .  c a v i t y , the most a n t e r i o r p o r t i o n of  The buccal c a v i t y i s not d i f f e r e n t i a t e d  from the pharynx which f o l l o w s i t but i s defined as that part of the stomodaeum on which are i n s e r t e d the second group of d i l a t o r muscles, t a k i n g t h e i r o r i g i n s on the clypeus  or on the  -78c l y p e a l area of the head, and having t h e i r i n s e r t i o n s a n t e r i o r to  the f r o n t a l ganglion and i t s connectives (Snodgrass).  pharynx merges i n t o the l o n g t u b u l a r oesophagus.  The  Near the  middle of the t h i r d t h o r a c i c segment the oesophagus enlarges g r a d u a l l y to form the crop.  In the specimen  from which t h i s  d e s c r i p t i o n was made (a nymph which would have moulted a female "brachypterous r e p r o d u c t i v e ) , the crop was  to form  one-half  the l e n g t h of the oesophagus and about f o u r times i t s breadth. At  i t s p o s t e r i o r end, the crop tapers to a diameter  greater than that of the oesophagus. at  The g i z z a r d i s l o c a t e d  the a n t e r i o r end of t h i s t a p e r i n g s e c t i o n .  forms the oesophageal  valve.  slightly  Posteriorly i t  Up to the p o i n t where the oesopha-  gus enlarges to form the crop, the d i g e s t i v e t r a c t was p l a c e d a l o n g the meson.  The  centrally  crop i t s e l f moves to the l e f t  of  the body. Mesent eron. P o s t e r i o r to the oesophageal v a l v e , the mid gives r i s e to f o u r g a s t r i c caeca.  intestine  Their diameter i s twice that  of  the oesophagus*  They are twice as l o n g as broad.  of  each v a r i e s s l i g h t l y , both i n l e n g t h and diameter.  The  size  The  extend as f a r forward as the a n t e r i o r end of the g i z z a r d .  caeca The  mid i n t e s t i n e passes from the g a s t r i c caeca, curving to the r i g h t and over the l a r g e i n t e s t i n e at a p o i n t between the vest i b u l e and the i n t e s t i n a l caeca and thence goes s l i g h t l y ward.  It  for-  v a r i e s i n width from three to f o u r times the diameter  of  the oesophagus, becoming s l i g h t l y "broader as i t approaches  the p o i n t of o r i g i n of the Malpighian the mid  to  At t h i s point  i n t e s t i n e i s v e n t r a l to the g a s t r i c caeca and  once more the alimentary enteron  tubules.  canal i s on the l e f t  side.  therefore The  mes-  extends t h e r e f o r e from the oesophageal v a l v e a n t e r i o r  the g a s t r i c caeca to the Malpighian  tubules.  Proctodaeum. The  slight  c o n s t r i c t i o n j u s t a n t e r i o r to the  i s the boundary between the mid todaeum.  tubules  and hind i n t e s t i n e , the proc-  There are ten Malpighian  tubules p a s s i n g p o s t e r i o r l y ,  t w i n i n g around the v a r i o u s p a r t s of the h i n d i n t e s t i n e  and  reaching as f a r back as the rectum.  smaller  at  They are s l i g h t l y  the p o i n t where they a r i s e from the hind i n t e s t i n e and  about o n e - f i f t h the diameter of the oesophagus*  are  P o s t e r i o r to  the tubules, the d i g e s t i v e t r a c t tapers to nearly one-half i t s o r i g i n a l diameter at the p y l o r i c or p o s t v e n t r i c u l a r valve„ region of the alimentary  canal i s the p o s t v e n t r i c u l u s and i s  about twice as l o n g as broad. tending to the l e f t , enlarged  This  P o s t e r i o r to the valve and  ex-  the h i n d i n t e s t i n e becomes enormously  to form the v e s t i b u l e ( C h i l d i n K o f o i d '34).  It i s  h e m i s p h e r i c a l i n o u t l i n e and opens a n t e r i o r l y , a f t e r a s l i g h t c o n s t r i c t i o n , i n t o the i n t e s t i n a l caecum, a s l i g h t p r o j e c t i o n of  the l a r g e i n t e s t i n e extending  to the l e f t .  The  slight  con-  s t r i c t i o n above mentioned i s v e n t r a l to the a n t e r i o r p a r t of the mes%teron.  From the i n t e s t i n a l caecum, the l a r g e i n t e s t i n e  -80goes forward n e a r l y and  as f a r as  as f a r as the a n t e r i o r  the s a l i v a r y r e s e r v o i r .  i s s i t u a t e d on the l e f t testine i s nearly v e n t r a l l y and mid  crop.  of the  crop  Here the proctodaeum At t h i s point  twice the diameter of the  crop.  the i n Passing  p o s t e r i o r l y , d o r s a l to the p o s t e r i o r end  i n t e s t i n e and  right  of the  end  v e n t r a l to the i n t e s t i n a l  caecum and  of  the  on  the  side of the v e s t i b u l e , the l a r g e i n t e s t i n e tapers "but  s l i g h t l y throughout i t s l e n g t h .  At a p o i n t midway alongside  the v e s t i b u l e , the proctodaeum tapers to a tube, whose diameter equals that of the g a s t r i c caeca, the l o n g as broad*  colon.  It r i s e s d o r s a l l y to point  This i s twice as  s l i g h t l y dorsal  to the r i g h t of the p o s t e r i o r margin of the v e s t i b u l e *  and  Here  i t becomes much enlarged, b e i n g about three times the diameter of the  colon,  to form the rectum which opens i n the anus.  The  rectum i s as long as broad. Accessary Glands of the Alimentary; Canal. The applied end  s a l i v a r y glands and  to the oesophagus and  of the  crop.  their reservoirs l i e closely  extend s l i g h t l y past the  Each gland c o n s i s t s of two  p a r t s , an  smaller  d o r s a l p o r t i o n and  surface  of the glands have a lumpy appearance due  rangement of the the d o r s a l and  anterior anterior  a posterior ventral portion.  c e l l s of which they c o n s i s t .  to the  The ar-  L y i n g between  v e n t r a l p a r t s of each gland i s a r e s e r v o i r .  i s about three times the diameter of the oesophagus and times as long as broad.  It reaches to the t i p of the  It  three  ventral  -81p o r t i o n of the gland.  Ducts from "both the glands and  reser-  v o i r s pass forward i n t o the head beneath the oesophagus.  Be-  tween the pharynx and the f l o o r of the head capsule, the gland duct and r e s e r v o i r duct of each side u n i t e to form a s i n g l e duct.  On approaching the base of the hypopharynx the two  ducts  become c l o s e l y a p p l i e d to one another j u s t before opening into the c a v i t y between the labium and hypopharynx. The mandibular  glands are,present around the pockets  between the mandibles and the m a x i l l a e . possess an e x t e r n a l opening.  They do not seem to  The glands are lbbed.  Zootermopsis  nevadensis.  The d i g e s t i v e system of t h i s species i s the same as that of the p r e c e d i n g s p e c i e s . Reticulitermes  hesperus.  As b e f o r e the alimentary canal opens by way b u c c a l c a v i t y which l e a d s to the pharynx.  of the  F o l l o w i n g t h i s the  oesophagus i s a l o n g tube extending to the crop which may may  or  not be d i f f e r e n t i a t e d e x t e r n a l l y , depending on the amount  of food which i t contains* about  When i t does contain food i t i s  three times the diameter of the oesophagus and  quite short, being s l i g h t l y longer than broad.  relatively  At the proven-  t r i c u l a r v a l v e a t the p o s t e r i o r end of the crop there i s a marked increase i n s i z e to form the g i z z a r d . h a l f times as l o n g as broad.  It i s one and  P o s t e r i o r l y It tapers to l e s s  than h a l f i t s o r i g i n a l diameter forming a short tube before  one-  -82merging i n t o the mid  i n t e s t i n e thus forming the oesophageal  v a l v e . * :, At the a n t e r i o r end  of the mid  i n t e s t i n e , the g a s t r i c  caeca are represented  by a c o l l a r extending s l i g h t l y a n t e r i o r l y  over the stomodaeum.  The mesenteron i s of the same diameter  as the g i z z a r d .  From the l e f t  s i d e , the mid  i n t e s t i n e passes  to the r i g h t on the d o r s a l s i d e , then curves v e n t r a l l y passing o b l i q u e l y forward to the l e f t abdominal segment. postventriculus. two  and  tapers  one-half  At  v e n t r a l side as f a r as the second  Here i t loops this point  going caudally to form the  the mid  i n t e s t i n e i s about  times the diameter of the oesophagus.  to a diameter s l i g h t l y greater  phagus a n t e r i o r to the Malpighian  It  than that of the oeso-  tubules.  A n t e r i o r l y the proctodaeum i s marked by a sudden increase  i n the s i z e of the d i g e s t i v e t r a c t and by the presence  of eight M a l p i g h i a n t u b u l e s . the l e f t  The p o s t v e n t r i c u l u s , s t i l l  v e n t r a l s i d e , moves c a u d a l l y , while t a p e r i n g  Hear the end  slightly.  of the f o u r t h abdominal segment i n the postven-  t r i c u l u s , i s the p y l o r i c v a l v e . an extensive  on  Immediately p o s t e r i o r there i s  enlargement to form the v e s t i b u l e and  the main  p a r t of the l a r g e i n t e s t i n e which moves to the r i g h t , d o r s a l l y beneath the mid  i n t e s t i n e , and much forward j u s t p o s t e r i o r to  the c a p h a l i c curve of the mesenteron.  At t h i s p o i n t i t begins  to taper as i t loops p o s t e r i o r l y coming to l i e p a r t l y on r i g h t s i d e of the enlarged  p o r t i o n of the hind i n t e s t i n e .  on the r i g h t s i d e the proctodaeum loops to the l e f t g i v i n g r i s e to the rectum.  the  before  Still  -83The s a l i v a r y glands are much l i k e those of Zootermopsis, They reach as f a r as the end of the f i r s t  abdominal segment  p o s t e r i o r l y and the second t h o r a c i c segment a n t e r i o r l y .  The  ducts of the glands open a n t e r i o r l y and separately i n a c h i t i n l i n e d tube, p r o b a b l y the s a l i v a r i u m . The mandibular glands c o n s i s t of simply-lobed bodies with a short duct opening on each s i d e of the pharynx between the  mandibles and m a x i l l a e .  THE REPRODUCTIVE SYSTEM. Female A l a t e . Zootermopsis a n g u s t i c o l i i s . As i n a l l i n s e c t s the female reproductive system cons i s t s of o v a r i e s , o v i d u c t s , vagina, c o l l e t e r i a l glands and a seminal r e c e p t a c l e . The o v a r i e s as a whole are f a s t e n e d a n t e r i o r l y by a f i n e t h r e a d - l i k e median ligament to the heart near the a n t e r i o r margin of the mesonotum where i t becomes fan-shaped.  This l i g a -  ment i s i n turn connected w i t h the a n t e r i o r t i p s of the terminal f i l a m e n t s of each o v a r i o l e at about the middle of the second abdominal segment.  The two  o v a r i e s l i e d o r s a l l y on the l a r g e  i n t e s t i n e , curving v e n t r a l l y around the d i g e s t i v e t r a c t i n the neighborhood of the f i f t h abdominal segment. s i s t s of between f o r t y and f i f t y type.  Each ovary con-  o v a r i o l e s of the p a n o i s t i c  The o v a r i o l e s pass v e n t r a l l y j o i n i n g the l a t e r a l  ducts independently and are arranged i n s e r i e s .  ovi-  The bases of the  o v a r i o l e s extend a l o n g the oviduct only as f a r as the middle of  -84the s i x t h abdominal s t e r n i t e .  The two l a t e r a l  oviducts,  mc r e a s i n g , i n s i z e as they pass p o s t e r i o r l y , f i n a l l y u n i t e to form the common oviduct abdominal s t e r n i t e .  at about the middle of the seventh  On i t s d o r s a l s u r f a c e , the median oviduct  r e c e i v e s the opening o f the seminal r e c e p t a c l e , an i r r e g u l a r l y lobed  s t r u c t u r e of about the same width as the median oviduct.  The r e c e p t a c l e i s a s a c - l i k e body. becomes the vagina.  Posteriorly this  oviduct  3?rom the n i n t h s t e r n i t e there a r i s e s a  median muscular tube, the median duct of the c o l l e t e r i a l which soon f o r k s d o r s a l l y .  glands,  The contents of these glands empty  i n t o the extreme p o s t e r i o r p o r t i o n of the median oviduct and p o s t e r i o r to the eighth s t e r n i t e .  Each f o r k bears on its d o r s a l  t i p a l a r g e number of convuluted tubes.  Those of the a n t e r i o r  f o r k a r e about the same s i z e as the Malpighian tubules;  those  of the p o s t e r i o r f o r k are much smaller i n diameter and also shorter.  The opening of the oviduct  i s l o c a t e d between the  e i g h t h s t e r n i t e and v e n t r a l v a l v u l a e .  The c o l l e t e r i a l glands  open i n the n i n t h s t e r n i t e p o s t e r i o r to the v e n t r a l v a l v u l a e ( P l a t e XVI, f i g s . 4 and 5). colleterial  Both the seminal r e c e p t a c l e and  glands l i e below the alimentary  connected to the r e s t of the reproductive  canal and are  system by the mediun  oviduct.  Zootermopsis nevadensis. This species has the female reproductive l a r to that of the preceding  species.  system sim-  -85Reticuliterxa es hesperus. In t h i s species the median ligament i s attached to the heart at the p o s t e r i o r margin of the metanotum.  The t i p s  of the o v a r i e s extend from the d o r s a l p a r t of the f o u r t h abdominal segment to the seventh s t e r n i t e , thus l o o p i n g the d i gestive t r a c t .  Each ovary c o n s i s t s of f i v e or s i x o v a r i o l e s .  At the seventh s t e r n i t e , the l a t e r a l o v i d u c t s from t h e i r l a t e r a l p o s i t i o n s turn m e d i a l l y , almost a t r i g h t angles, at the same time becoming reduced i n s i z e . to form a short median o v i d u c t .  The two ducts u n i t e  Just p o s t e r i o r to t h i s duct  i s a.seminal r e c e p t a c l e , smallest at i t s opening and broadly rounded at i t s t i p . The c o l l e t e r i a l glands are almost  identical  with those of Zooterrnopsis i n that the common duct i s twobranched and muscular.  The tubules are fewer i n number and  much s h o r t e r than i n Zooterrnopsis. They a r e about twice the diameter of the Malpighian t u b u l e s .  The openings of these  three above d e s c r i b e d organs a r e s i m i l a r l y placed as i n the two p r e c e d i n g s p e c i e s , a l l opening into the median o v i d u c t . Brachypterous and Apterous Reproductives. Zooterrnopsis a n g u s t i c o i l i s . The female r e p r o d u c t i v e system o f these castes a r e l i k e those of the a l a t e  caste.  In R e t i c u l i t e r m e s f l a v i p e s , the female sex organs are smaller than those of the a l a t e s , those o f apterous forms being the s m a l l e s t .  • -86Soldiers. ' •  Zootermopsis a n g u s t i c o l i i s . In t h i s caste the median ligament  i s very  long  spreading out fan-shaped a t i t s a n t e r i o r connection.  The  o v a r i e s themselves extend from the t h i r d abdominal segment posteriorly*  Those few d i s s e c t e d showed considerable v a r i a t i o n  i n the development of the o v a r i e s .  There are at l e a s t twenty  o v a r i o l e s i n each ovary with the eggs only s l i g h t l y developed. I have not found the same degree of a r r e s t e d development of the ova and the misplacement of the n u c l e i which Thompson ('22) records.  As i n the other castes the l a t e r a l oviducts pass  v e n t r a l l y i n the r e g i o n of the seventh abdominal segment. However, they a r e considerably much elongated above the intersegmental  extending  from  membrane to the median oviduct  sit-  uated p o s t e r i o r l y to the l a s t abdominal ganglion.  This duct  i s quite short, tapers sharply caudally jpid i t s caudal p o r t i o n A.  p r a c t i c a l l y covered by the seminal  vesicle.  The r e c e p t a c l e i s  a f l a t f o u r - l o b e d s t r u c t u r e d o r s a l to the union l a t e r a l ducts.  of the two  In one specimen examined the r e c e p t a c l e was  connected to the median o v i d u c t . i n some s o l d i e r s the seminal  Thompson ('22) states that  r e c e p t a c l e i s not connected with  the o v i d u c t . Reticulitermes. In s o l d i e r s of R. flavip.es, the seminal  receptacle  and the c o l l e t e r i a l glands a r e not connected with the median oviduct  (Thompson and Synder '20).  developed i n s o l d i e r s .  The organs a r e l e a s t  -87Male M a t e . *  Zooterrnopsis  nevadensis.  In a l l i n s e c t s , the r e p r o d u c t i v e  system c o n s i s t s of  a p a i r of t e s t e s , two vasa d e f e r e n t i a , a median e j a c u l a t o r y duct and sometimes a p a i r of seminal v e s i c l e s . In t h i s s p e c i e s ,  the two t e s t e s are nearly s p h e r i c a l ,  fmm. i n skiameter or about o n e - t h i r d  the width of the pronotum,  the surface not u n l i k e that of mulberries t i p s of the sperm tubes.  The l e f t  p o s t e r i o r to the v e s t i b u l e *  t e s t i s l i e s a short"distan  Conducting the spermatozoa f r  the t e s t e s are two l o n g slender about one and one-half  due to the rounded  om  vasa d e f e r e n t i a , i n length  times the diameter o f the t e s t i s .  are about the same diameter as the Malpighian tubules.  They They  pass v e n t r a l l y c l o s e to the v e n t r a l abdominal w a l l , l a t e r i n g d o r s a l l y to pour t h e i r contents separately antero-dorsal  ce  w a l l of the e j a c u l a t o r y duct.  vasa d e f e r e n t i a become l a r g e r u n t i l  loop-  through the  Near the duct the  they are of t h e i r l a r g e s t  diameter where they u n i t e with the e j a c u l a t o r y duct.  Immediate-  l y a n t e r i o r to the openings of the vasa d e f e r e n t i a are the separate openings of the ducts of the accessory  glands.  These  ducts a r e stout, about twice as long as broad, and a n t e r i o r l y possess about s i x much branched tubules, the diameter o f . t h e Malpighian tubules ducts of the glands.  each nearly three  times  and twice as long as the  The e j a c u l a t o r y duct i s broadest at the  openings of the vasa d e f e r e n t i a , being h a l f the width of the t e s t i s .  s l i g h t l y l e s s than one-  At t h i s p o i n t , the e j a c u l a t o r y  -88cluct v e n t r a l l y i s s l i g h t l y sory grand ducts. the p e n i s .  two-lobed,  extending below the acces-  The duct tapers sharply forming the t i p of  The whole s t r u c t u r e of the e j a c u l a t o r y duct i s  s t r o n g l y muscular, i t s lumen b e i n g l i m i t e d c o n s i d e r a b l y by dings of the inner w a l l .  fol-  This copulatory mechanism l i e s pos-  t e r i o r to the l a s t abdominal g a n g l i o n and opens at the p o s t e r i o r margin of the n i n t h s t e r n i t e , where the p o s t e r i o r t w o - f i f t h s are surrounded by a c a v i t y between the n i n t h and tenth s t e r n i t e s . The spermatozoa are much elongated. Discussion:  In the w r i t i n g s of Thompson ('22), I rams  ( ' 24) and K i r b y ('34), the accessory glands above described are termed the seminal v e s i c l e s . glandular i n function.  However, the tubules are d e f i n i t e l y  Thompson r e a l i z e d t h i s and even stated  that no spermatozoa were found i n the tubules of any caste or age.  Sperms may  be found a l l along the vasa d e f e r e n t i a .  seminal v e s i c l e s are, s t r i c t l y vasa d e f e r e n t i a (Snodgrass)  Since  speaking, only d i l a t i o n s of the  there i s l i t t l e doubt as to the  absence of such storage organs.  The v e n t r a l b u l b u l a r nature of  the e j a c u l a t o r y duct i s not a modified seminal v e s i c l e . Zootermopsis  angusticoliis.  The specimen of t h i s s p e c i e s used was l y the same age as the above.  or about  approximate-  However, the t e s t e s , although  s i m i l a r l y p l a c e d , were much smaller, being about diameter  of  4/lOmra. i n  o n e - f i f t h the width of the pronotum;  testis has about a dozen sperm tubes.  The  The vasa def e r e n t i a , i n  length, are twice the width of the t e s t i s .  The duct of the  -89left  accessory gland was broader  than the r i g h t , the l a t t e r  being l i k e that i n the preceding s p e c i e s . had In  Some small tubules  t h e i r source at the base of the two ducts of the glands. other r e s p e c t s the system i s the same. R e t i c u l i t e r m e s hesperus * The system of t h i s s p e c i e s 1B a l i t t l e  different.  There a r e fewer sperm tubes, u s u a l l y between f i v e and ten and r e l a t i v e l y they a r e l a r g e r when compared to the whole t e s t i s . 4 ,* u", . -' The l e f t t e s t i s i s ventral^lro the rectum, the r i g h t . Opposite the intersegmental membrane. more v e n t r a l . diameter  The l e f t  i s therefore s l i g h t l y  Both a r e l o c a t e d i n the seventh segment.  The  of the vasa d e f e r e n t i a i s the same as that of the  Malpighian tubules and are as l o n g as f i v e times the width of the t e s t e s .  Leaving the t e s t e s , they pass v e n t r a l l y  to the  n i n t h s t e r n i t e and m e d i a l l y to the d o r s a l surface of the ejacul a t o r y duct with which they u n i t e c e n t r a l l y * course, the l e f t  vas deferens i s p r a c t i c a l l y v e n t r a l .  and v e n t r a l to the openings separate openings glands.  Throughout i t s Anterior  of the seminal ducts are the two  of the ducts of the two l o n g - c o i l e d accessory  Each gland i s as wide as one-half the diameter  t e s t e s and n e a r l y as long as the vasa d e f e r e n t i a .  of the  P o s t e r i o r to  the f o u r ducts i s the broad e j a c u l a t o r y duct, i t s width greater than the o u t s i d e d i s t a n c e between the two anal  slightly stylets.  P o s t e r i o r l y i t tapers to form the p e n i s which opens at the caudal tip  of the n i n t h s t e r n i t e .  -90Brachypterous Reproductives. gootermopsis  angusticoliis.  The t e s t e s are much l a r g e r and c o n s i s t of many more sperm tubes*  Each t e s t i s i s about 1mm.  one-half the width of the pronotum.  i n diameter and n e a r l y  At the a n t e r i o r t i p s of the  ducts of the accessory glands, there are more less-branched tubules, u s u a l l y about a doaen. and u n i t e a l o n g t h e i r b o r d e r s .  Some of these bend on themselve The e j a c u l a t o r y duct i s not as  broad as i n the a l a t e s but t h i s may be j u s t a v a r i a t i o n .  In  other r e s p e c t s the system i s much l i k e that of a l a t e s . Reticulitermes. R; f l a v i p e s possesses l a r g e r t e s t e s than-alates (Thompson and Snyder '20). Apterous Reproductives. Zootermopsis  angusticoliis.  This caste has the t e s t e s developed to a greater extent than any other c a s t e ; tubes i n each  There a r e at l e a s t f i f t y  sperm  testis. R e t i c u l i t ermes.  The same i s true f o r t h i s species as f o r the precedi n g although the sperm tube number i s not the same. Soldiers. Zoot ermopsis In the adult  angusticoliis•  s o l d i e r the t e s t e s are l a r g e r than i n the  -91a l a t e s , although rauch smaller than those of the "brachypterous and apterous r e p r o d u c t i v e s , "but the vasa d e f e r e n t i a are slender and c o n t r a c t e d , almost without lumen, and the "seminal v e s i c l e s " are v e s t i g i a l i n s t r u c t u r e (Thompson *22). lobes a r e broad and rounded,  Few  of the test.es-'  as i n the r e p r o d u c t i v e castes, but  are l o n g and u s u a l l y s l e n d e r , s e v e r a l small lobes merging  into  one l a r g e r l o b e , and f i n a l l y a l l u n i t i n g at the base of the t e s t i s i n t o one common space, with which the vas deferens i s connected.  The p a i r e d c h a r a c t e r of the "seminal v e s i c l e s " i s  e s p e c i a l l y w e l l seen i n the s o l d i e r  (Thompson).  Thompson states  that owing to the degeneration of p a r t s of the system,  sper-  matids and tubules of the glands, the male s o l d i e r i s probably s t e r i l e as i s the case with the females. Reticulitermes. *  fla-vipes the t e s t e s are small, the vasa deferen-  n  t i a s t r a i g h t and the "seminal v e s i c l e s " are not convoluted (Thompson '20). probably  An embryonic  p e n i s i s present.  T h i s caste i s  sterile. Immature. Stages. Zootermopsis Sex organs may  nymphs.(Thompson  '19).  angusticollis_.  be d i s t i n g u i s h e d i n newly hatched  -92THE NERVOUS SYSTEM. Th'e 'Central Nervous System. Alates. Zootermopsis a n g u s t i c o l i i s . As  i n the m a j o r i t y of i n s e c t s the c e n t r a l nervous  system c o n s i s t s p r i n c i p a l l y of the b r a i n , a suboesophageal ganglion,  three t h o r a c i c g a n g l i a and a variable number of ab-  dominal g a n g l i a . The b r a i n of a l a t e termites n e a r l y f i l l s width of the head, extending  the e n t i r e  from the b i f u r c a t i o n of the e p i -  c r a n i a l arms to n e a r l y h a l f the d i s t a n c e to the f r o n t o - c l y p e a l suture.  In the b r a i n , the protocerebrum i s by f a r the l a r g e s t  p a r t of the b r a i n * lobes extending  Viewed a n t e r o - d o r s a l l y i t appears as two  c a u d a l l y and a n t e r i o r l y . L a t e r a l l y and s l i g h t -  l y v e n t r a l , each p r o t o c e r e b r a l lobe gives r i s e to the o p t i c lobes.  These are about one-half  of the p r o t o c e r e b r a l l o b e s . head capsule,  the l e n g t h ( a n t e r o - p o s t e r i o r l y )  T r a v e r s i n g the Bpace across the  the lobes become s l i g h t l y  c o n s t r i c t e d and s p l i t  into three p a r t s , the middle part b e i n g one-half of the other  two.  more and spread  the diameter  As they approach the eye they u n i t e once  fan-shaped over the inner surface of the eye.  Ventrad and a n t e r o - l a t e r a d of the p r o t o c e r e b r a l lobes are the deutocerebral  lobes.  These are much smaller than the o p t i c  lobes and give r i s e to the antennal deutocerebral  nerves.  V e n t r a l to the  lobes are the lobes of the tritocerebrum which  pass around the d i g e s t i v e t r a c t .  They taper p o s t e r o - v e n t r a l l y .  -93On t h e i r inner margins they are u n i t e d "by the small cerebral•commissure.  trite  V e n t r a l to the commissure are the t  quite l a r g e circumoesophageal of the t r i t o c e r e b r a l l o b e s .  1  connectives, v e n t r a l extensions A r i s i n g from the a n t e r i o r surface  of these lobes and dorsad of the t r i t o c e r e b r a l commissure are the f r o n t o - l a b r a l nerves.  From each of these a r i s e two  nerves,  the outer supplying the labrum and f r o n t , the inner one passing to the f r o n t a l ganglion (to be d e s c r i b e d under the stomodaeal nervous  system). The f r o n t a l gland, above and  caudal to the main  p o r t i o n of the b r a i n , ^mentioned by^Thompson ('22);is much d o u b t , f u n c t i o n l e s s . it  The  without  question might a r i s e as to whether  i s even a. fnnt-ion-l-e-s-s gland.  It i s true that c e l l s at this  p o i n t are d i f f e r e n t from the other hypodermal c e l l s of the head^in b e i n g much elongated.  However, at t h e i r v e n t r a l mar-  gins are the o r i g i n s of muscles whose i n s e r t i o n s are i n the oesophagus. At the a n t e r i o r end of the v e n t r a l nerve suboesophageal ganglion. laterally,  cord i s the  I t i s roughly ovoid i n shape.  Viewed  the connectives u n i t e with i t p o s t e r o - d o r s a l l y .  On  the upper a n t e r i o r s u r f a c e there are a p a i r of mandibular nerves, c l o s e together at the ganglion but V e n t r a l to t h i s p a i r of nerves  soon separating widely.  i s another widely  separated  p a i r , the m a x i l l / a r y nerves.  Each of these has a branch  i n g i n t o the hypopharynx.  the a n t e r o - v e n t r a l surface of  On  the ganglion i s the p a i r of widely  separated l a b i a l  pass-  nerves.  -94At the v e n t r a l and p o s t e r i o r t i p are the p a i r e d connectives which pass v e n t r a l l y beneath the tentorium to the thorax. m a x i l l a r y nerves a r e s l i g h t l y l a r g e r i n diameter l a b i a l nerves. •fd-Jt&citf^pass The  The  than the  From the m a x i l l a r y and l a b i a l nerves  branches  i n t o the p a l p i . t h o r a c i c g a n g l i a are s i t u a t e d a n t e r i o r to the  coxal c a v i t i e s .  From the suboesophageal ganglion with the  g a n g l i a of the head.  A l l three t h o r a c i c g a n g l i a a r e e s s e n t i a l l y  the same, i n c r e a s i n g i n s i z e from the prothorax to raetathorax, and a l l have three p a i r s of nerves.  The proganglion and meta-  g a n g l i o n are about the same shape, being n e a r l y c i r c u l a r . second  t h o r a c i c ganglion tapers a n t e r i o r l y .  that i n the prothorax the f i r s t  Child  The  ('34) s t a t e s  p a i r of nerves are small c o r -  1 responding to the nerves s u p p l y i n g the wings i n the other thoracic  segments*  In s e c t i o n e d m a t e r i a l (8th i n s t a r ,  r e p r o d u c t i v e ) there was found  to be a p a i r of l a r g e  fertile  nerves  d p a s s i n g forward and l a t e r a l l y , probably to the c e r v i c a l r e g i o n . These nerves a r e as l a r g e as the nerves of the l e g s which are the l a r g e s t nerves l e a v i n g the t h o r a c i c g a n g l i a . thorax and raetathorax the f i r s t  In the meso-  p a i r or wing nerves pass  for-  ward a l o n g the connectives f o r a short d i s t a n c e then diverge q u i c k l y going d o r s a l l y of  to supply the wings.  The second  v e i n s s u p p l i e s the t h o r a c i c muscles and w a l l .  In the same  specimen above mentioned these were small nerves j u s t to  the l e g nerves i n the prothorax.  pair  anterior  In the other two segments  they a r i s e from about the middle of the g a n g l i a and pass obl i q u e l y forward*  They are l a r g e r than the wing nerves.  -95From the l a t e r o - c a u d a l end of each g a n g l i a i s a p a i r of l a r g nerves-passing v e n t r a l l y to each p a i r of l e g s .  e  They have a  small "branch not f a r d i s t a n t from the ganglion which supplie the thorax*  The  s connectives to and away from the g a n g l i a are  l a r g e r than the l e g nerves and  except f o r the a n t e r i o r con-  n e c t i v e s of the raetaganglion<> each.pair l i e s c l o s e together;-  Y The a n t e r i o r connectives of t h i s ganglion are widely In two  separated.  i n d i v i d u a l s examined the a n t e r i o r connectives of the  p r o t h o r a c i c g a n g l i a were twisted one over the other.  The  g a n g l i a and connectives l i e v e n t r a ^ to the muscles fastened to the spinae.  -9.6-  There are s i x abdominal -segments, one i n each of the abdominal segments^, two to seven.  The f i r s t , f i v e are a l l small g a n g l i a .  Each has two p a i r s of nerves p a s s i n g l a t e r a l l y ^ r o m the ..ganglia) to  the v e n t r a l abdominal w a l l , l y i n g beneath the muscles  which  l i n e the f l o o r of the abdomen and supply^ the s p i r a c l e s as well as the muscles of the segments. to  the f i r s t  first  The s i x t h ganglion i n a d d i t i o n  two p a i r of n e r v e s h a s three other p a i r s . ?  of these i s very s m a l l .  The second i s d i s t i n c t l y  and much branched, s u p p l y i n g the eighth segment.  The larger  The t h i r d and  by f a r the l a r g e s t of these or any other nerve of the abdomen passes around the r e p r o d u c t i v e organs, g i v i n g o f f to them at l e a s t one p a i r of nerves, and s u p p l i e s the n i n t h and tenth segments where there are branches to the muscles, anus and (in  stylets  the case of males). Zooterrnopsis nevadensis. The c e n t r a l nervous system i s l i k e that of the pre-  ceding s p e c i e s . R e t i c u l i t e r m e s hesperus. The b r a i n i s e s s e n t i a l l y the same as the two above d e s c r i b e d .  species  However, there are a few important d i f f e r e n c e s ,  namely two o c e l l a r nerves l o c a t e d on the a n t e r i o r aspect of the b r a i n and a f u n c t i o n a l f r o n t a l gland with a f o n t a n e l nerve.  As  viewed d o r s a l l y t h e p o s t e r i o r margin of the b r a i n i s nearly..; s t r a i g h t and i s emarginate c e n t r a l l y where the f r o n t a l gland i s situated. the  brain.  rounded  The f o n t a n e l nerve may  be seen p a s s i n g a n t e r i o r l y to  A n t e r i o r l y the b r a i n becomes broader towards the  optic  ../  -97lobes.  Instead of a s i n g l e o p t i c nerve there are many f i n e  t h r e a d - l i k e nerves.  The d e u t o c e r e b r a l lobes are r e l a t i v e l y  s l i g h t l y l a r g e r than i n Zootermopsis. The suboesophageal g a n g l i o n i s of the same shape as i n Zootermopsis and has three p a i r s of nerves. Of the t h o r a c i c ganglia,, that of the metathorax i s the  l a r g e s t while the p r o t h o r a c i c one i s the smallest.  Each  has three p a i r s of nerves, the l e g nerves having a small branch going to the thorax. There a r e s i x abdominal g a n g l i a l o c a t e d i n the second to  seventh segments.  JS^s^-^he  first  of these g a n g l i a i s  n e a r l y as l a r g e as the p r o t h o r a c i c ganglion but is, s p i n d l e shaped and not ovoid. largest  The s i x t h abdominal ganglion i s the  of th.c-crtircra-and i s "slightly l a r g e r than the f i r s t .  I have not been able to examine the abdominal nerves except those of the s i x t h ganglion*  These nerves supply the eighth  and n i n t h segments, the t i p of the abdomen, and the reproduct i v e organs. Brachypterous Reproductives. Zootermopsis a u g u s t i c o l l i s . In  t h i s caste the o p t i c l o b e s of the b r a i n are  s l i g h t l y reduced.  There a r e two o p t i c nerves i n s t e a d of three  as i n the a l a t e s .  A c c o r d i n g to Thompson ('22), the f r o n t a l  gland i s absent. i s the same. the  The remainder of the c e n t r a l nervous system  The g a n g l i a of the thorax and abdomen are much  same as i n a l a t e s .  -98Apt erous Reproductives. Zoot ermopsis  angusticoliis.  The "brain i s smaller than that of the a l a t e , on account of the r e d u c t i o n of the o p t i c lobes (Thompson '22). Soldiers. Zoot ermopsis a n g u s t i c o l i i s . The "brain of the s o l d i e r i s smaller than that of the a l a t e or apterous r e p r o d u c t i v e "but i s of the same s i z e as the brachypterous r e p r o d u c t i v e . The g a n g l i a of the thorax and abdomen a r e l i k e those of a l a t e s . Reticuldtermes hesperus. As i n the a l a t e s , the f i r s t abdominal ganglion i s l a r g e r than the next f o u r . Immature Stages. Zootermopsis  angusticoliis.  The castes have been separated i n newly hatched nymphs by the s i z e of the b r a i n (Thompson '19). The p r i n c i p a l p a r t s of the b r a i n may "be d i s t i n g u i s h e d . The Stomodaeal ITervo-us System. Zootermopsis  angusticoliis.  The system, sometimes c a l l e d t h e " a n t e r i o r sympathetic nervous system^consists of the f r o n t a l ganglion s i t u a t e d on the d o r s a l w a l l of the oesophagus, two connectives from the t r i t o c e r e b r a l l o b e s and^fecurrent nerve p a s s i n g caudally, d o r s a l to  -99the  oesophagus and v e n t r a l to the "brain.  The loop formed "by  the  f r o n t a l ganglion connectives i s almost c i r c u l a r .  The  ganglion i s about two-thirds the diameter of' the circumoesophageal connectives i n c r o s s - s e c t i o n and i s longer than broad. It  i s s i t u a t e d a n t e r i o r to the b r a i n .  The recurrent nerve i s  of  small diameter b e i n g about a f i f t h  of  the g a n g l i o n as seen i n v e r t i c a l l o n g i t u d i n a l s e c t i o n .  of the small dimension A  small nerve passes a n t e r i o r l y s u p p l y i n g the c l y p e a l region. R e t i c u l i t e r m e s hesperus. A small f r o n t a l ganglion appears c r a d l e - l i k e i n cross-section.  The remainder of the system has not been  studied. THE 'SENSE ORGANS. Zootermopsis a n g u s t i c o i l i s . In  t e r m i t e s there a r e three types of p e r i p h e r a l sense  organs, namely, the organ of Johnston, sensory pores and sensory h a i r s . The f i r s t  of these, the organ of Johnston, i s the  l a r g e s t , and i s l o c a t e d on the second antennal segment.  The  c e l l s and nerves are arranged i n an i n v e r t e d cone and terminate in  the membrane between the second and t h i r d antennal segments.  Other sense organs of the antennae consist of pores near the a n t e r i o r margin of the second segment (Noyes '30). of  On the rest  the antennae are sensory h a i r s s u p p l i e d with t i n y nerves.  -99A-  A second nerve center of the stomodaeal system i s a p a i r of g a n g l i a connected at t h e i r p o s t e r i o r margins to form a l a r g e median o c c i p i t a l ganglion ©sESs-situated beneath the a o r t a . T h i s ganglion connects w i t h a nerve running d o r s a l to the oesophagus - j^robably the r e c u r r e n t nerve from the frontsil ganglion*  The  o c c i p i t a l ganglion has  two  large  connectives  u n i t i n g with the v e n t r a l side of the b r a i n between the oesophageal  connectives.  (to be i n s e r t e d on Page 99, a f t e r l i n e  8)  circum  -leoOn the labrum Noyes has found no sensory pores. my  In  i n v e s t i g a t i o n s I have observed pores or p i t s on the t i p of  the labrum of newly hatched nymphs.  On the v e n t r a l side of  the clypeus j u s t p o s t e r i o r to the l a b r o - c l y p e a l suture there are  s i x groups of c i r c u l a r pores, each group having from about  15 to 30 In each.  In newly hatched t e r m i t e s the pores are  l i m i t e d to two groups of a few pores i n each.  Each group i s  p l a c e d l a t e r a d of the epipharynx. In and l a c i n a .  the m a x i l l a e pores are found on the p a l p , galea The pores of the palp are l o c a t e d on the v e n t r a l  s u r f a c e s of the f i r s t  three segments. pro  D o r s a l l y the galea has  mad  a group of pores j u s t ji-o-g-terier to i t s t i p . There are pores a l s o on the v e n t r a l s u r f a c e . of  Near the t i p of the v e n t r a l tooth  the l a c i n a there i s a group  of pores*  Sensory h a i r s are  present on both s u r f a c e s of the p a r t s of the m a x i l l a e . The pores of the labium are c o n f i n e d to the glossae and p a r a g l o s s a e .  As i n the m a x i l l a e , sensory h a i r s are  s c a t t e r e d over a l l the p a r t s of the labium. There i s a group  of pores s i t u a t e d on the p o s t e r i o r  suspensoria of the hypopharynx. The most prominent  sense organ of the legs i s s i t -  uated on the outer s u r f a c e of the t i b i a e near the proximal ;  end. of cal  At t h i s p o i n t  very small pores.  the organ i s marked e x t e r n a l l y by a group I n t e r n a l l y these pores l e a d to a s p h e r i -  body c o n s i s t i n g of many c e l l s .  The h i s t o l o g i c a l  details  and the exact nature of the organ have not yet been worked out. Noyes has found that the area i s w e l l s u p p l i e d w i t h nerve  -101endings.  I have "been a b l e to l o c a t e t h i s organ on a l l three  p a i r s of legs of a l l castes and ages f o r "both B r i t i s h  Columbia  s p e c i e s of Zootermopsis, R e t i c u l i t e r m e s hesperus and Cop tot ermes f O rmO SanUS  r d m Shanghai-,-**"  Of the other p a r t s of the l e g s , pores are found only on the t r o c h a n t e r .  These occur i n three groups, one on the  inner s u r f a c e and two on the outer s u r f a c e .  Sensory  hairs  are p r e s e n t , however, on the other p a r t s of the l e g s . Sensory h a i r s are present on the c e r c i and abdominal styles. THE TRACHEAL SYSTEM* Zootermopsis  angusticoliis  As i n a l l i n s e c t s the t r a c h e a l system c o n s i s t s of a s e r i e s of l o n g i t u d i n a l , d o r s a l and v e n t r a l t r a c h e a l trunks a r i s i n g from a main trunk which has i t s o r i g i n i n the s p i r a c l e s . In t e r m i t e s there are two t h o r a c i c s p i r a c l e s and eight abdominal spiracles.  The two s p i r a c l e s of the thorax are, s t r i c t l y  speaking, those of the l a s t  two t h o r a c i c segments although  .they are found i n the membranes between the pro and mesothorax and the meso and metathorax.  These two s p i r a c l e s are oval i n  shape and are longest d o r s o - v e n t r a l l y . openings a r e two l i p s , h a l f the opening.  Extending over the  the a n t e r i o r one covering more than  The p o s t e r i o r l i p or p o s t l a b i a i s l e s s  than one-half the width of the p r e l a b i a and i s much l e s s sclerotized.  The atrium i s l i n e d with a network of s n a i l  l i k e structures.  The a t r i u m i s immediately  separated into  cup-  -102two  t r a c h e a l trunks. " ', Of the abdominal s p i r a c l e s the f i r s t  largest.  i s by f a r the  Sumner ('33), although not making a d e t a i l e d  study  of t h i s s p i r a c l e , suggested that i t might p o s s i b l y be homologous w i t h the tympanum of the A c r i d i i d a e . I have been unable to l o c a t e any typanum.  gitn---excuaim-t-3^m^ The s p i r a c l e i t -  s e l f i s c i r c u l a r i n o u t l i n e and without l a b i a e .  There i s ,  however, a r i n g o f c h i t i n l e s s e n i n g the diameter of the aperture.  I n t e r n a l l y the a t r i u m broadens to n e a r l y twice i t s  diameter w i t h the s i d e s l i n e d with the same network described f o r the t h o r a c i c s p i r a c l e s . t r a c h e a l trunk.  Here i t g i v e s r i s e t e a l a r g e  The other abdominal s p i r a c l e s are considerably  much s m a l l e r and occur on a l l the segments to the e i g h t h .  =103ORGAITS OF DISTRIBUTION,. CONSERVATION AND The  ELIMINATION.  Blood. Zooterrnopsis a n g u s t i c o i l i s .  Of the "blood, only the c e l l u l a r p a r t has been amined.  The  blood  shaped, b e i n g  corpuscles  or haemocytes are roughly  three times as broad as the;; are deep.  the diameter across  Nucleolus can be d i s t i n q u i s h e d . u l e s are q u i t e d i s t i n c t . and  The  nu-  the c e l l .  A  Chromatin net knots or gran-  The blood moves forward i n the  heart  aorta* The The  Organs of C i r c u l a t i o n .  d o r s a l blood  v e s s e l i s the only part of  haemocoele c l o s e d by d e f i n i t e w a l l s . an a n t e r i o r a o r t a and  the  It c o n s i s t s of two  a p o s t e r i o r heart.  The heart  a n t e r i o r margin of the mesonotum. i n the l a s t  two  There are eleven  t h o r a c i c segments.  broadest j u s t a n t e r i o r to the  intersegmental  a n t e r i o r are l o c a t e d a p a i r of s l i t - l i k e side.  A n t e r i o r l y the  The  chambers taper  the metathorax i s the l a r g e s t and t h i r d the width of the chamber.  parts,  extends  d o r s a l l y .from the p o s t e r i o r t i p of the n i n t h t e r g i t e to  two  disc-  From the  t h i n edge of the c e l l s there are short pseudopodia. c l e i extend nearly one-half  ex-  the  chambers,  chambers are membrane.  o s t i a , one  slightly.  The  Slightly  on each chamber of  tapers to a tube nearly The  one-  chamber of the mesothorax  tapers to a tube t u r n i n g s l i g h t l y v e n t r a l through the thorax. On  e n t e r i n g the prothorax t h i s tube or a o r t a enlarges  slightly  -104d moving to a p o i n t j u s t dorsa^l <bfr the oesophagus i n the c e r v i c a l r e g i o n . " i n the head the a o r t a gus, to the o c c i p i t a l ganglion.  continues dorsad 66 the oesophaHere the v e n t r a l w a l l disappears  The  d o r s a l wall becomes very broad extending to beneath the b r a i n  and  l i m i t e d by the a n t e r i o r nerves of the o c c i p i t a l The  ganglion.  number of beats i s v a r i a b l e depending on the i n -  s t a r and the time i n that p e r i o d .  In a s i x t h i n s t a r  there  were t h i r t y - t h r e e beats per minute. THE EAT BODY. Zoot ermopsis a n g u s t i c o l l i s . The  f a t body c o n s i s t s of l a r g e c e l l s grouped into  v a r i o u s p o l y g o n a l shapes and i s found c h i e f l y i n the abdomen and  d o r s a l to the oesophagus i n the thorax.  the f a t body i s f l a t  In the abdomen  extending over the d i g e s t i v e t r a c t .  THE DEH00YTES. Zootermopsis a n g u s t i c o l i i s . The  oenocytes I have not i d e n t i f i e d w i t h c e r t a i n t y .  Snodgrass s t a t e s that ded  the oenocytes of Isoptera  remain embed-  i n the epidermis a t t h e i r p o i n t s of o r i g i n and extend as  segmental c l u s t e r s i n t o the body c a v i t y . THE CORPORA ALATA. d The  corpora a l a t a l i e almost v e n t r a l pB the caudal  p o r t i o n of the b r a i n i n c l o s e contact They a r e more or l e s s oval i n shape.  with the oesophagus.  PART I I BIONOMICS LIFE HISTORY. The C o l o n i z i n g F l i g h t , Swarmirg and PairjLjng. The c o l o n i z i n g f l i g h t  i s f o r the purpose of found-  i n g new c o l o n i e s and -age-not to be confused w i t h , n u p t i a l With  f l i g h t s of other i n s e c t s , the f l i g h t  -is" t e r m i t e s , p r e c e d i n g the time f o r  to appear, a p e r t u r e s are opened i n the wood to  allow the emergence of the a l a t e s .  This u s u a l l y occurs during  the months of August, September and October f o r Zooterrnopsis  T  b e i n g dependent  on temperature and moisture c o n d i t i o n s of the  l a t e summer evenings.  R e t i c u l i t e r m e s hesperus does not appear  until after a r a i n f a l l  so that the c o l o n i z i n g f l i g h t may be  delayed u n t i l the winter months.  However, a l a t e s of a l l three  B r i t i s h Columbia species may be seen f l y i n g during p r a c t i c a l l y all  the months of the year.  Guarding the exit holes are s o l -  d i e r s w i t h t h e i r mandibles p o i n t i n g outwards.  At the c r u c i a l  moment l a r g e niambers of a l a t e s w i l l pass out i n t o the open and then take f l i g h t .  The wing beat of Zooterrnopsis i s slow but  powerful, that of R. hesperus weak. towards higher l i g h t  Usually the former f l i e s  i n t e n s i t y and may not come to rest f o r  s e v e r a l hundred yards.  R. hesperus being a weak f l i e r i s  u s u a l l y p i c k e d up by the wind and i s c a r r i e d  considerable  distances. E v e n t u a l l y the swarms s e t t l e down to the ground near, or on wood i n the case of the damp-wood t e r m i t e s .  Almost  immediately they shed t h e i r wings by moving backwards-, cat ching  -106t h e i r w i n g s on p r o j e c t i n g p a r t s , t h e n m o v i n g s i d e w a y s . ually  t h i s t a k e s hut a matter o f minutes.  The w e s t e r n  Ussubter-  ranean t e r m i t e b r e a k s o f f i t s wings by r a i s i n g the t i p of the abdomen*  The f e m a l e s o f a l l  t h r e e species at t h i s time r a i s e  t h e t i p s o f t h e i r abdomen, p r o b a b l y e m i t t i n g an odor to t h e raales^and a w a i t t h e a r r i v a l .of m a t e s .  After  t h e f e m a l e l e a d s t h e way w i t h t h e m a l e f o l l o w i n g ^ f o r a s u i t a b l e colony. in  s e a r c h o f wood.  attractive pairing  searching  Reticul^i_t,e.rme_s may e n t e r t h e g r o u n d  Z o o t e r m o p s i s i s r e s t r i c t e d t o wood  lying  to  on t h e g r o u n d , o r ^ s t u m p s .  When s u i t a b l e wood i s f o u n d a c a v i t y  i s made, o n l y by t h e f e m a l e o f Z o o t e r m o p s i s b u t by b o t h s e x e s i n R. h e s p e r u s * the p a i r  A s soon a s t h e c a v i t y i s l a r g e enough t o h o l d  i t s o p e n i n g i s s e a l e d w i t h p e l l e t s , a n d c h i p s o f wood  sealed together with l i q u i d  feces.  Pert i l i z a t i o n . I n t h e c a s e o f R e t i c u l i t e r m e s c o p u l a t i o n may o c c u r t h e day f o l l o w i n g b u t i n Z o o t e r m o p s i s i t does n o t u s u a l l y o c c u r . until  two weeks h a v e p a s s e d .  C o p u l a t i o n i s a c c o m p l i s h e d by  •the m a l e a n d f e m a l e p l a c i n g t h e t i p s o f t h e i r abdomens t o g e t h e r a n d r e m a i n i n g i n t h i s p o s i t i o n f r o m one t o f o u r m i n u t e s . Egg-laying. U s u a l l y R. h e s p e r u s l a y s on t h e a v e r a g e eggs w i t h t h e l a s t f e w l a i d a t i n c r e a s i n g l y l o n g Eggs are' - f i r s t s , l a i d } In 1  c o l o n i e s of Zootermopsis  u s u a l l y , , i n t h e t h i r d week.  about t e n intervals.  angusticoliis  An a v e r a g e o f a b o u t t w e l v e c o m p l e t e s  -107the f i r s t  deposition.  F o l l o w i n g t h i s no more eggs are l a i d  u n t i l the f o l l o w i n g s p r i n g .  In l a b o r a t o r y  c o l o n i e s of Z.  nevadensis no eggs had appeared a f t e r the s e l e c t i o n of mates, (^after seven months*  T h i s may be due to unnatural  conditions  but Z. a n g u s t i c o l l i s i n the l a b o r a t o r y under the same conditions agreed quite c l o s e l y to the above statements. egg p r o d u c t i o n  The r a t e of  v a r i e s with the type of r e p r o d u c t i v e .  Third  form or apterous r e p r o d u c t i v e s may l a y as many as f o r t y eggs i n two days. Care of Eggs. The eggs a r e u s u a l l y p l a c e d to r e c e i v e the optimum temperature and humidity.  They may be c a r r i e d from one p l a c e  to another by the a d u l t s who h o l d them between t h e i r mandibles. In a d d i t i o n the eggs a r e r e g u l a r l y cleaned by the f i r s t in  form  the case of new c o l o n i e s , but by nymphs i n o l d c o l o n i e s . Incubation  Period.  In Z. a n g u s t i c o i l i s the incubation p e r i o d v a r i e s between f o r t y - f o u r and e i g h t y - e i g h t days with an average of fifty  s i x days.  In one of my experiments eggs l a i d by sup-  plementary r e p r o d u c t i v e s i n s t a r nymphs-,within  hatched when cared f o r by seventh  s i x t y - s e v e n days.  For R. hesperus the  p e r i o d v a r i e s between t h i r t y and one hundred days,  averaging  about f i f t y  first,  to f i f t y - s i x days.  U s u a l l y , eggs l a i d  hatch i n a s h o r t e r l e n g t h of time.  -108! B i o t i c \ P— o— t— e n—t— i— a— l, i ———-j As  i n a l l t e r m i t e s ^ t h e j b i o t i c j p o t e n t i a l of females  i s u s u a l l y q u i t e high. 2500-3000.  For R.  emerged females.  For Z. a n g u s t i c o i l i s i t i s  approximately  hesperus i t i s about one hundred f o r newly This f i g u r e i s probably  very  conservative.  Developmental Stages. Embryoni c• As the eggs contain a l a r g e amount of yolk with the cytoplasmic m a t e r i a l ^ d i s t r i b u t e d around the periphery the egg and about the nucleus, At the end of the tenth day  the cleavage  of  i s meroblastic.  there i s developed a c a p - l i k e  s t r u c t u r e at the l a r g e end of the egg.  Within  days t h i s g i v e s r i s e to the embryonic head.  the next ten  Prom i t and  ex-  tending down the outside of the convex surface i s the embryo. First  signs of segmentation appear followed by  of the appendages.  the  At t h i s time the embryo extends  the d i s t a n c e to the other end of the egg.  of the small end of the egg.  one-half  At the end of twenty-  f i v e days the embryo extends to about one-third the  ger and  beginnings  distance  The head appendages are much l o n -  are d i s t i n c t l y d i f f e r e n t i a t e d from the much smaller  t h o r a c i c appendages.  The  abdomen i s represented  up k n o t - l i k e s t r u c t u r e at the p o s t e r i o r end.  by a c u r l e d  At the end of  t h i r t y - f i v e days the t h o r a c i c appendages have become quite long and  extend n e a r l y to the small end of the egg.  domen i s more developed having  reached the end of the  The  ab-  egg.  -109At  t h i s time the embryo has begun to encroach quite n o t i c e a b l y  on the center and body of the y o l k mass.  At f o r t y days (old)  the p l e u r a of the embryo are b e g i n n i n g to become pronounced • and by the end of the f o r t y - f i f t h day the body -wall has practically  enclosed the remaining p o r t i o n of the y o l k .  p o i n t the abdominal  At  this  c e r c i are w e l l d i f f e r e n t i a t e d and the  segmentation of the abdomen i s complete.  The remaining time  r e q u i r e d f o r h a t c h i n g (about twenty days), i s spent, i n the completion of the development  of the i n t e r n a l  structures.  Po st embryonic. Young fixa.t-inutar-^ymphs of Z. a i i g u s t i c o l l i s emerge from the egg by an opening which i s s p l i t  three ways, one  ex-  tending down on each s i d e of the head, the t h i r d along the d o r s a l m'eson.  The nymph i s pure white except f o r the t i p s of  the mandibles and some of the spines of the l e g s , which are light  brown.  stomodaeally  During the ten days of t h i s stage i t i s fed (from s a l i v a r y  glands?) by the a d u l t s or nymphs.  In R i hesperus the young nymph i s u s u a l l y from i t s s h e l l .  assisted  I t s food c o n s i s t s of m a t e r i a l taken from the  mouth or anus of o l d e r nymphs or a d u l t s or of brown f e c a l plaster.  The f i r s t  instar  i s passed i n about f o u r t e e n to  twenty days.*? F o l l o w i n g the fi.rs..t i n s t a r p e r i o d the yo\mg nymph moults e n t e r i n g the second p-e^i-ed—O-F- i n s t a r . the nymphs are f e d p r o c t o d a e a l l y .  During t h i s p e r i o d  This p e r i o d i s twenty  f o r Z. a n g u s t i c o l i i s and about a month f o r R. hesperus•  days  -110In R. hesperus the t h i r d i n s t a r r e q u i r e s a month and the f o u r t h in  two months.  the s i x t h i n s t a r .  sixth instar.  Dwarf workers appear i n young c o l o n i e s  Reproductive nymphs a l s o appear i n the  In o l d e r c o l o n i e s the l a r g e r workers are i n the  seventh i n s t a r .  A l a t e s a t t a i n f u l l development i n the seventh  instar. In Z. a n g u s t i c o i l i s the t h i r d i n s t a r r e q u i r e s from twenty-two  to twenty-four days, the f o u r t h about f o r t y - f i v e  days and f o r p s e u d o - s o l d i e r nymphs, twenty days i n the f i f t h . These s o l d i e r nymphs moult to become adult s i x t h instax".  s o l d i e r s i n the  In o l d e r c o l o n i e s s o l d i e r s may become mature  as l a t e as the e i g h t h or n i n t h i n s t a r s .  The f i r s t  p r o d u c t i v e nymph which moults to become an a l a t e seventh  form r e -  i s in i t s  instar.  P o s t - a d u l t Gr_ow_th and Longevity. In a l a t e females of Z. a n g u s t i c o i l i s post-adult growth i s apparent by the general appearance of the abdomen. The segments of the abdomen become separated, the i n f o l d i n g s -between each segment becoming l e s s deep. membranes between to t h e i r f u l l e s t larger  The intersegmental  the t e r g i t e s and s t e r n i t e s a r e y a l m o s V s t r e t c h e d ^ extent.  Although the male adult  does become  than the newly emerged a l a t e i t s p o s t - a d u l t growth i s  much l e s s than that  of females.  This i s due i n the case of  females to an i n c r e a s e i n the number of egg tubes and also to an a c t u a l  c e l l u l a r d i v i s i o n and growth.  l i n g or d i s t e n s i o n  It i s not a mere swel-  (Snyder 3 5 ) . The p o s t - a d u l t growth of males 1  -Illi s due t o c e l l u l a r  increase.  T h i s may he "brought a b o u t b y t h e  s p e c i a l -stomodaeal f o o d r e c e i v e d f r o m nymphs. ©  m o n t h s =>W  I n my c o l o n i e s o f L . n ey a _ d e n s i s ^ t h e r e a r e y e t no broods and p o s t - a d u l t  growth i s ' c o n s i d e r a b l y l e s s  pronounced.  T h e r e i s an i n c r e a s e i n t h e s i z e o f t h e abdomen i n b o t h s e x e s , s i n c e t h e segments a r e more c l e a r l y d i f f e r e n t i a t e d n e w l y emerged In a d u l t growth. at  least  than i n  alates. R. h e s p e r u s , t h e f e m a l e s undergo an enormous p o s t The t h o r a x becomes much b r o a d e r a n d t h e abdomen  t h r e e t i m e s t h e l e n g t h o f t h e abdomen o f a newly  a l a t e female.  emerged  The s c l e r i t e s o f t h e abdomen a p p e a r a s s m a l l  s l e n d e r c u r v e d b a r s w i d e l y s e p a r a t e d by membrane.  I n s u c h queens  •there i s a d e g e n e r a t i o n o f t h e wing muscles and t h e muscles o f the  j a w s s i n c e t h e y no l o n g e r f e e d t h e m s e l v e s b u t a r e f e d by t h e  w o r k e r s a n d r e p r o d u c t i v e nymphs. do n o t show much p o s t - a d u l t  The m a l e s o f t h i s  species  growth.  The a d u l t s o f A . a n g u s t i c o i l i s may l i v e  about  five  years i R o l e o f S u p p l ementary Rep_roduct i v e s . The a p t e r o u s o r b r a c h y p t e r o u s s u p p l e m e n t a r y  repro-  d u c t i v e s a p p e a r when e i t h e r o f t h e r o y a l p a i r d i e o r i f t h e y become s e p a r a t e d f r o m t h e r e s t o f t h e c o l o n y due t o an i n c r e a s e in i t s size.  The r o y a l p a i r  t h e n h a v e no c o n t r o l l i n g  o v e r t h e most d i s t a n t p a r t s o f t h e c o l o n y .  influence  When s u c h a c o n d i t i o n  r e s u l t s i n a c o l o n y , c e r t a i n nymphs, u s u a l l y t h o s e i n t h e f o u r t h to  s e v e n t h i n s t a r s , b e g i n t o undergo a development  not l i k e  that  -112of younger nymphs. reproductives. mature.  These a r e the developing  supplementary  Soon they become straw-coloured  and sexually  Should one of the r o y a l p a i r d i e then that one i s r e -  placed "by a supplementary r e p r o d u c t i v e of the same sex.  Should  both d i e or "become separated  then  from the r e s t of the colony  supplementary r e p r o d u c t i v e s of "both sexes w i l l appear. the immortality of the colony ficient  food m a t e r i a l  And so  i s made p o s s i b l e , p r o v i d i n g suf-  exists.  These supplementary r e p r o d u c t i v e s have l a r g e r reprod u c t i v e organs than a l a t e s or the r o y a l p a i r .  The supplementary  queens may l a y as many eggs i n one day as the r o y a l queen does in two y e a r s .  The t e s t e s of s u p p l e m e n t a l reproductives of  Z. a n g u s t i c o l i i s are nearly twice the s i z e of a l a t e t e s t e s . Therefore with the i n t r o d u c t i o n of supplementary forms into a colony  i t s r a t e of growth i s i n c r e a s e d tremendously.  The  C o n s t i t u t i o n o f the Colony. The  castes of any one colony vary with  year and the age of the colony. a y e a r - o l d colony w i l l  the time of  In the case of Z. a n g u s t i c o l i i s ,  c o n s i s t of the r o y a l p a i r , one s o l d i e r  and from twelve to twenty nymphs.  With an increase i n the age  of c o l o n i e s there appear more s o l d i e r s i n the r a t i o of s o l d i e r s to nymphs as one to f i f t y .  During  the greater part of the year  only adult r e p r o d u c t i v e s , s o l d i e r s and nymphs are present colony.  i n the  With the approach of the l a t e summer months there begin  to appear l a r g e numbers of the a l a t e caste.  C o r r e l a t e d with the  appearance of the a l a t e s i s the increase i n number of s o l d i e r s .  -113Immediately f o l l o w i n g swarming of the a l a t e s there i s a drop i n the percentage of s o l d i e r s .  They g r a d u a l l y "become k i l l e d  a c e r t a i n p o i n t which i s apparently  stable.  Soon i n the o l d  c o l o n i e s , that i s , i n the f a l l months, the t i n y f i r s t nymphs appear i n f a r greater numbers than at any the  o f f to  instar  other time of  year. Only i n o l d c o l o n i e s w i l l any,supplementary reproduc-  t i v e s be found. been recorded  T h e i r number v a r i e s considerably but  i t has  that a colony headed by such queens and kings  c o n s i s t i n g of about two  thousand i n d i v i d u a l s may  supplementary r e p r o d u c t i v e s . quite c o n s e r v a t i v e . are no other a d u l t  My  There may  have  and  twenty-six  opinion i s that t h i s f i g u r e i s  be f e r t i l e s o l d i e r s where there  reproductives.  D i s t r i b u t i o n w i t h i n the Colony. The  d i s t r i b u t i o n of the termites w i t h i n the colony i s  subject to change during the d i f f e r e n t O r d i n a r i l y these humidity ally treat  changes are due  seasons of the year.  to unfavorable  temperature or  c o n d i t i o n s or to the amount of food a v a i l a b l e .  Gener-  the colony d u r i n g the winter months w i l l ^ i f possible-^reto a p o i n t below the f r o s t  case where the colony to at l e a s t  l i n e of the ground, or i n the  i s l o c a t e d i n huge stumps, the r e t r e a t i s  the inner recesses of the colony.  With the approach  of warmer weather there w i l l be a r e t u r n to warmer quarters. When excessive d e s s i e a t i o n occurs owing to the l a c k of any c i p i t a t i o n over long p e r i o d s R.  hesperus w i l l probably  pre-  l i v e in  the deepest p a r t s of the colony where i t i s able to get i t s  -114necessary  moisture.  As a colony  grows older c e r t a i n p a r t s of i t  become deprived of food so that the termites have to move to other p a r t s of the (colony. Our  damp-wood s p e c i e s , Zooterrnopsis. a r e very u n l i k e  those of many of the t r o p i c a l forms i n which the r o y a l p a i r are confined to a c e l l .  In some of these  t r o p i c a l forms there are  a l a r g e number of termites tending the queen as w e l l as a l a r g e number of s o l d i e r s .  In Zootermopsis, the r o y a l p a i r moves about  quite f r e e l y with the r e s t of the colony. w i t h i n the colony,  Of a l l the forms  only the eggs have any d e f i n i t e l o c a t i o n  w i t h i n the colony and even t h i s i s subject to change*  But,, the  l o c a t i o n i s the same with r e f e r e n c e to optimum temperature and humidity  c o n d i t i o n s , since the eggs are moved to a p l a c e  having  optimum c o n d i t i o n s whenever there i s a change of such c o n d i t i o n s . Theory of Caste D i f f e r e n t i a t i o n , Having thus studied the l i f e h i s t o r y and the castes of the t e r m i t e B i t i s now time to i n v e s t i g a t e the d i f f e r e n t i a t i o n of these  castes, that i s , how they came to be. Before d i s c u s s i n g t h i s a b r i e f examination of some  i n s e c t s of the order I'HyWenoptera i n which there i s caste d i f f e r e n t i a t i o n w i l l help  considerably i n understanding  the problem.  In the genus Meiipona the c e l l s i n which the males, queens and workers are reared a r e a l l a l i k e and of the same s i z e (Wheeler '28).  They are each given the same kind of food and an egg i s  l a i d i n each a f t e r which the c e l l s are sealed. treatment r e s u l t s i n the production  The uniform  of two sharply  differentiated  -115feraale castes, the workers and  queens.  The  queen has  i n g very* small o v a r i e s with immature eggs. Trigona the queen c e l l s are l a r g e r and quantity of food. r i p e eggs. was  The  It.would  on emerg- •  In the a l l i e d genus  c o n s i s t of a greater  queen on emerging has her o v a r i e s f u l l of  seem then that the p r o v i s i o n of more food  a stimulus i n a c c e l e r a t i n g the development of her o v a r i e s .  Much the same i s t r u e f o r the honey "bee i n which the queen r e ceives " r o y a l j e l l y "  throughout her l a r v a l existence and  the wor-  kers /-this food f o r a p e r i o d of only four days followed by a food of lower q u a l i t y * f i v e days before  In t h i s case the queen emerges four to  the worker with her o v a r i e s nearly mature as  against the s l i g h t l y developed o v a r i e s of the worker. i n s e c t s , the caste d i f f e r e n t i a t i o n of females i s due  In to  these  environ-  mental or e x t r i n s i c f a c t o r s . Returning we  to t e r m i t e s , i n p a r t i c u l a r to Z. a n g u s t i c o l i i s ,  f i n d that the caste d i f f e r e n t i a t i o n occurs  Also there seeas to be evidence  i n both sexes.  that some castes a r i s e due  to  f a c t o r s e x i s t i n g i n the germ plasm , that i s , i n t r i n s i c f a c t o r s . Other castes seem to be d i f f e r e n t i a t e d by enviromental f a c t o r s . It does not seem u n l i k e l y that s o l d i e r s are t i a t e d long before  environmental f a c t o r s could have any  on t h e i r development. Emmeraon c l e a r l y ted by two  differen-  The observations  effect  of Imms, Heath and  i n d i c a t e that each sex was  originally  represen-  f e r t i l e forms, a winged small-headed form and an  terous or brachypterous large-headed  s o l d i e r , which was  ap-  differen-  t i a t e d from the small-headed type and l a t e r became s t e r i l e .  -116Further, the s o l d i e r Is thought not to "be derived from workers. Supporti-ng  these statements are the f o l l o w i n g f a c t s : .  s o l d i e r s are sometimes f e r t i l e or approach f e r t i l i t y Zooterrnopsis.  2. Some s o l d i e r s possess wing "buds.  1. P r i m i t i v e as i n  How  could  s o l d i e r s with wing buds be d e r i v e d from apterous forms? 3. P r i m i t i v e s o l d i e r s u s u a l l y have compound eyes developed to some degree and present.  i n some cases  the v e s t i g e s of l a t e r a l o c e l l i  4. S o l d i e r s are always present  are l a c k i n g * i s the f i f t h  In i n c i p i e n t  e v o l u t i o n a r y development?  even where the workers  c o l o n i e s the f i r s t  instar soldier.  are  adult to appear  Is t h i s a r e c a p i t u l a t i o n of i t s  However, as w i l l be shown l a t e r ,  c e r t a i n f e a t u r e s of the development of s o l d i e r s can only be explained by The evidence  environmental or e x t r i n s i c f a c t o r s . experimental  to the theory  work of C a s t l e ( 34)  concrete  e x p l a i n i n g caste d i f f e r e n t i a t i o n by  t r i n s i c or environmental f a c t o r s . theory  gives  $  He  c a l l s i t the  ex-  inhibition  s i n c e , i n a study of r e p r o d u c t i v e forms, the development  of a r e p r o d u c t i v e of any members of that sex. become separated  sex i s governed by  the presence of  In other words should a group of nymphs  from f u n c t i o n a l r e p r o d u c t i v e s , f u n c t i o n a l straw-  colored supplementary r e p r o d u c t i v e s of both sexes w i l l w i t h i n a p e r i o d of t h i r t y - f i v e to f i f t y  days.  In the presence  of a f u n c t i o n a l r e p r o d u c t i v e of a given sex,' no r e p r o d u c t i v e s of that sex develop* ment i s t r u e . d u c t i v e s may  C a s t l e has  The  develop  supplementary  converse of t h i s s t a t e -  found that the supplementary repro-  be e i t h e r apterous or brachypterous.  The  former  -117are derived from nymphs i n the f o u r t h to s e v e n t h . i n s t a r s . l a t t e r a r e developed from nymphs i n the f i f t h  to seventh  The instars.  In a d d i t i o n he has found that a l l members of a colony are potentially  r e p r o d u c t i v e and any i n d i v i d u a l ,  except the pseudo-  s o l d i e r nymph, the adult s o l d i e r , the brachypterous nymph l a t e in  i t s seventh i n s t a r and soon to become a l a t e and the f i r s t  form or a l a t e r e p r o d u c t i v e s , may become a supplementary reprod u c t i v e i n the absence of a f u n c t i o n a l r e p r o d u c t i v e of i t s sex. An a l c o h o l or an ether e x t r a c t prepared  from the bodies of  f u n c t i o n a l queens when f e d to nymphs m a t e r i a l l y increases the time f o r the development of female supplementary but  i t does not a f f e c t  productives  reproductives  the development of supplementary r e -  of the opposite.sex.  From these conclusions, i t  would seem that from the f u n c t i o n a l r e p r o d u c t i v e s the nymphs obtain some substance, p o s s i b l y a s e c r e t i o n or could i t be that proctodaeal  food d e r i v e d from the f u n c t i o n a l reproductives by  the nymphs i s e f f e c t i v e i n i n h i b i t i n g the development of t h e i r r e p r o d u c t i v e organs?  Is i t p o s s i b l e that the f e c a l m a t e r i a l of  f u n c t i o n a l r e p r o d u c t i v e s i s d i f f e r e n t from that of other forms to  the extent  to be proven.  that i t i s i n h i b i t o r y i n nature?  This remains  In any case the presence of the f u n c t i o n a l r e -  productive i s s u f f i c i e n t  to. deter the development of the r e -  p r o d u c t i v e organs of the nymphs.  This would then seem to be a  case of environmental c o n d i t i o n b r i n g i n g about caste d i f f e r e n tiation. Experimenting  with s o l d i e r s , C a s t l e has found some  -118rather  i n t e r e s t i n g r e l a t i o n s h i p s o f s o l d i e r s w i t h nymphs w h i c h . t o  p o i n t p a r t l y e x t r i n s i c t h e o r i e s and p a r t l y to g e n e t i c o f i n t r i n sic an  theories. incipient  As was s t a t e d above t h e f i r s t colony  i s a fifth-instar  t h e nymphs a r e i n t h e f i r s t , soldier  appear.  This  times, but always w i t h y o u n g nymphs t h e r e nymphs o f t h e f i f t h is ony  g r e a t l y delayed.  soldier.  At t h i s  second o r t h i i d i n s t a r s .  i s removed f r o m s u c h a c o l o n y  dier w i l l  a d u l t to appear i n  another f i f t h  e x p e r i m e n t may be r e p e a t e d the same'results.  i s placed  instar  with the r o y a l p a i r  t h e appearance of s o l d i e r s into a col-  a s o l d i e r o r s o l d i e r nymph, t h e  appearance of another s o l d i e r i s prevented or g r e a t l y Apparently  sol-  several  S h o u l d a s o l d i e r be i n t r o d u c e d  not a l r e a d y p o s s e s s i n g  I f the  I f i n s t e a d of the  i n the colony  and s i x t h i n s t a r s ,  time  the presence of s o l d i e r s i s i n h i b i t o r y  delayed.  on t h e d e v e l -  opment o f more s o l d i e r s w h i c h i s a l m o s t t h e same as was t r u e f o r supplementary r e p r o d u c t i v e s . why t h e s o l d i e r i s t h e f i r s t  However t h i s does n o t e x p l a i n adult  t o a p p e a r i n an i n c i p i e n t  colony. In c o n c l u s i o n ,  i t may be s t a t e d t h a t up t o t h e p r e s e n t  t i m e many t h e o r i e s h a v e b e e n p o s t u l a t e d b u t o n l y has  r e c e i v e d any e x p e r i m e n t a l  proof  that of Castle's  i n explaining the d i f f e r e n -  t i a t i o n of castes of termites. TROPISMS. Tropisms a r e forms of behaviour r e s u l t i n g from They a r e c l a s s i f i e d a c c o r d i n g The  tropisms  stimuli.  t o t h e s t i m u l i w h i c h p r o d u c e them.  of termites are h e l i o t r o p i s m ,  thermotropism,  -119hydrotropism, chemotropism. and • ; One  thigmotropisia.  of the most ob ious of the tropisms i s h e l i o t r o p i s m v  or the r e a c t i o n to l i g h t  stimuli*  At only one time d u r i n g t h e i r  l i f e h i s t o r y are any of the castes p o s i t i v e l y h e l i o t r o p i c .  At  swarming time a l a t e s are p o s i t i v e l y h e l i o t r o p i c , even f l y i n g i n t o areas of higher l i g h t  intensity.  It i s strange that  mediately f o l l o w i n g t h i s p e r i o d they "become n e g a t i v e l y tropic.  im-  helio-  A l l other castes and ages respond n e g a t i v e l y to l i g h t  hut the degree of r e a c t i o n i s dependent on the l i g h t  intensity  p r e v i o u s l y experienced "by them. Thermotropism  i s apparent only a f t e r experimental  s t u d i e s and c a r e f u l o b s e r v a t i o n s . and c o l d temperatures.  Termites d i s l i k e both hot  During the winter months they r e t r e a t  to the deeper recesses of the colony. found c l o s e r to the surface*  In any  case, the termites of  the damp-wood species w i l l be found  only  tunnels, having r e t r e a t e d from the d r i e r p a r t s .  i n the moist The f a c t  the eggs are c a r e f u l l y placed to r e c e i v e the optima of and humidity may  be  T h i s t r o p i c behaviour i s probably  c l o s e l y l i n k e d with hydrotropism. at l e a s t  In summer they may  that  temperature  be a t t r i b u t e d to a combination of both thermo-  tropism and hydrotropism. Thigmotropism  i s evident to a l l who  shedding of the wings which f o l l o w s swarming.  have observed the Almost  immediately  a f t e r the wings have been ruptured from t h e i r bases, the a l a t e s w i l l be seen to be h u r r y i n g about  searching f o r a p i e c e of wood  under which to crawl or a crack i n which to make a colony.  -120This  i s due  to t h e i r p o s i t i v e t h i g m o t r o p i c  I n t h e -tunnels  of the  some t h i g m o t r o p i c colony  the  colony  p a c k e d i n t o each  t h e nymphs and  behaviour *  t e r m i t e s w i l l be  show  opening a  tunnel.  experimental  f l a v i p e s was  castes  o b s e r v e d to b e , e x t r e m e l y w e l l  as y e t  not  received  w o r k t o make i t c o n c l u s i v e t h a t  are n e g a t i v e l y chemotropic. £•  other  A l m o s t i n v a r i a b l y on  C h e m o t r o p i s m i n t e r m i t e s has sufficient  characteristics.  negatively  Hartwell  ( 24) 1  has  shown  they  that  chemotropic to c e r t a i n e s s e n t i a l  oils. HABITS. Nearly pect any  to t h e i r  a l l termites are very  t u n n e l s but  also i n respect  g r o u p i s c a r e f u l l y w a t c h e d , one  tennae are  carefully  and palpus..  cleaned  F o r k e g s may  o f t h e body a r e l i c k e d t e r m i t e to be  be  t i d y , not  c l e a n by  i n the  same way.  groomed r e m a i n s q u i e t w h i l e  i t s groomer  the  an-  mandibles  palps in  The carefiilly the  cleaning. i s that of  In t h e p r o c e s s o f g r o o m i n g an appendage may  to eat  If  Other p a r t s  " r e c i p r o c a l grooming."  A h a b i t w h i c h commonly o c c u r s  which the  t h a t the  drawn b e t w e e n t h e  p a s s e s o v e r i t s body u s i n g i t s m a n d i b l e s and p r o c e s s of  in res-  to t h e i r b o d i e s .  w i l l perceive  by b e i n g  cleaned  only  groomer f e e d s on injured insect.  t e r m i t e s are  eaten.  the  exuding blood  Frequently  be  cannibalism.  cut o f f f o l l o w i n g  and  nay  weak, d i s e a s e d  proceed o r dead  -121ITATURAL ENEMIES. Our  n a t i v e termite s p e c i e s have no p a r a s i t e s and  only a few p r e d a t o r s . found  Guests or i n q u i l i n e s may  sometimes be  i n the c o l o n i e s . P r e d a t o r s are the most important  control*  animals  i n natural  Of these, evening f l y i n g b i r d s account f o r l a r g e  numbers of the a l a t e s during the c o l o n i z i n g f l i g h t . as woodpeckers have been observed  B i r d s such  d r i l l i n g into termite c o l o n i e s .  Amongst the i n v e r t e b r a t e s , p r e d a t o r s on t e r m i t e s at the time of swarming are  s p i d e r s , wasps and f l i e s .  other stages are  ants (Hymenoptera).  The  Predators of the smaller species  seem to a t t a c k the damp-wood termites more r e a d i l y than l a r g e r b l a c k carpenter ant.  T h i s ant i s often found  the  i n the  same stump with the tunnels of each i n s e c t i n t e r l o c k i n g and at times even connecting with one Mites and Z. a n g u s t i c o i l i s .  Collembola  another. are common i n the tunnels of  Mites are more f r e q u e n t l y found  to the bodies of the termites and the tunnel i t s e l f . l a t t e r were observed t e r m i t e bodies. not yet proven.  the Collembola  clinging  r e s t r i c t e d to  In some of the laboratory c o l o n i e s , the i n l a r g e numbers, even crawling on  Whether these two  the  animals are i n q u i l i n e s i s  PAR? I l l INTESTINAL EAUNA INTRODUCTION. Protozoa ly,  occur  the Orthoptera,  i n about s i x orders of i n s e c t s , name-  I s o p t e r a , Neuroptera,  doptera and D i p t e r a (Kudo '3l) . these orders^ except n e a r l y a l l belonging  The protozoa occurring i n vail  I s o p t e r a or termites.^are p a r a s i t i c species, to the C l a s s Sporozoa.  c e r t a i n species of mosquitos are found of trypanosomes. tera.  Coleoptera^, L e p i -  In D i p t e r a , i n  the i n v e r t e b r a t e stages  Sporozoa occur i n s e v e r a l species of Orthop-  In the wood roach, (Crypto cere us p u n c t u l a t u s J there are  non f r e e - l i v i n g protozoa  c l o s e l y resembling  those of termites  (Snyder '35). In the I s o p t e r a , both p a r a s i t i c and symbiotic species occur, b e l o n g i n g to the f o u r c l a s s e s of protozoa. Those of the termites of B r i t i s h Columbia a r e confined to two of these c l a s s e d , the Mastigophora ( f l a g e l l a t e s ) and the Sporozoa ( g r e g a r i n e s ) . In a d d i t i o n to protozoa o c c u r r i n g i n termites there are a l s o b a c t e r i a and s p i r o c h a e t e s .  In t h i s t h e s i s only the  protozoa w i l l be d e s c r i b e d . THE PROTOZOA Classification. Before proceeding list  to t h e i r  classification I shall  them a c c o r d i n g to the termite species i n which they 1.  Zooterrnopsis a n g u s t i c o i l i s and nevadensis Trichonympha c o l l a r ! s  Kirby.  occur:  -123T, • ;  campanula  T, sphaerica  K o f o i d and Swezy. ( K o f o i d and Swezy) Duboscq and Grasse.  Streblomastix s t r i x  Kofoid'and  Trichomonas termopsidis  Cleveland.  T r i c e r c o m i t u s termopsidis Hexamastix- termopsidis Hirmocystis t e r m i t i s K o f o i d i n a ovata 2.  Swezy.  Kirby.  Kirby. (Leidy)  Henry.  R e t i c u l i t e r m e s hesperus Trichonympha a g i l i s  Leidy.  Spironyrnpha  Koidzumi.  S.  porteri  ovalis  Sp i r o t r i c ho ny rap ha f l a g e l l a t a Microjoenia r a t c l i f f e i Torquenympha p_ct.pp.luB  Brown. Brown.  Holomast i g g t e s elongaturn Dinenympha f i m b r i a t a Pyrsonympha ma.i or £• minor  Kirby.  Powell.  Powell.  Hirmocyst i s t e r m i t i s These species arranged  (Leidy). a c c o r d i n g to t h e i r p o s i t i o n  t a b l e s (Kudo) are as f o l l o w s :  PHYLUM PROTOZOA Class Mast <j)gophora Subclass  Grassi.  Powell.  P. granulata  i n the c l a s s i f i c a t i o n  G r a s s i and Foa.  Zoomastigina  -124Order  Polymastigida  - •, T r i b e Mono20a Group 1.  Streblomastix  strix  Group 2.  Dinenympha f i m b r i a t a Pyrsonympha major P. minor P. g r a n u l a t a  Group 3.  Hexamastix termopsidis T r i c e r c o m i t u s termopsidis  Group 4. Order  Trichomonas termopaidis  Hypermastigida.  Family 1.  Holomastigotidae Holomastigotes  elongaturn  Spirotrichonympha  flagellata  Spironympha p o r t e r i S. Family 2.  ovalis.  Joenidae Microjoenia r a t c l i f f e i Torquenympha octoplus  Family 3. Trichonymphidae Trichonympha T. campanula T» s p h a e r i c a T. a g i l i s Class Sporozoa Subclass  Telosporidia  collaris  -125Order - '  ;  Gregarinida  Suborder  Eugregarinina  Legion  Cephalina  ^Hirmocystis I  termitis  Kofoidina  ovata.  These w i l l now be d e s c r i b e d order  i n which they appear i n t h i s  i n the accompanying  table.  i n the  They may be compared  illustrations.  Streblomastix having  i n greater d e t a i l  strix.  This i s an elongate  species  four f l a g e l l a l o c a t e d at the h o l d f a s t at the t i p of the  rostrum.  The nucleus i s an elongated  which c l e a r l y  spindle.  show are s p i r a l l y arranged.  The rayonemes  I t s length varies  c o n s i d e r a b l y , 15 - 320 microns. Dinenympha f i m b r i a t a . with four to eight f l a g e l l a r The  This species i s spindle-shaped  cords arranged i n s p i r a l form.  a x o s t y l e i s a rod, not always attached p o s t e r i o r l y .  It i s  about 60 microns i n l e n g t h . Pyrsonympha major.  T'his i s t y p i c a l l y  club-shaped,  t a p e r i n g uniformly, from the rounded p o s t e r i o r end to the more or l e s s p o i n t e d a n t e r i o r end. present terior  i n the endoplasm.  A few small granules  The a x o s t y l e i s attached  end.to the c e ^ t r o b l e p h l a r o p l a s t .  into two, three, or f o u r threads. flagellar  cords.  at i t s an-  It i s split posteriorly  There may be four to eight  The average l e n g t h i s about 110 microns.  Pyrsonympha granulata. a l a r g e number of densely i s 83 microns.  may be  The endoplasm u s u a l l y  s t a i n i n g granules.  contains  I t s average length  -126Pyrsonympha minor.  The endoplasm i s vacuolated.  The  average l e n g t h i s about 60 microns. Hexamastix t e r m o p s i d i s .  In shape t h i s t i n y  i s o v o i d a l or p y r i f o r m , and narrowed p o s t e r i o r l y . f i v e a n t e r i o r flagella^sometimes microns l o n g . the nucleus  species  There are  p r o x i m a l l y u n i t e d 1 5 to 25 ?  The a x o s t y l e i s slender, curves  and p r o j e c t s f o r a short d i s t a n c e .  c l o s e l y around The l e n g t h  v a r i e s between 5 and 11 microns. T r i c e r c o m i t u s termopsidis.  There are three a n t e r i o r  f l a g e l l a 6-20 microns l o n g and a very long t r a i l i n g f l a g e l l u m (19-65 microns long) which adheres to the body surface c l o s e to the p o s t e r i o r end. The body of the protozoa  The a x o s t y l e i s very  until  indistinct.  i s about 7 microns long.  Trichomonas t e r m o p s i d i s . c i r c u l a r but may be elongated * 7  This species i s u s u a l l y nearly  I t averages 43 microns i n length.  There a r e four a n t e r i o r f l a g e l l a about the l e n g t h of the body. The u n d u l a t i n g membrane i s l a r g e with f r e e f l a g e l l u m of moderate A  length*  The a x o s t y l e may be enlarged  minating  i n a short f i l a m e n t . Holomastigotes elongatura.  i n a plumb-bob form t e r -  The body i s spindle-shaped  and about 100 microns long.  F l a g e l l a are i n many c l o s e l y a r -  ranged s p i r a l s .  i s near the a n t e r i o r end.  The nucleus  Sp i r o t r i cho nympha f l a g e l l a t a .  The body i s about 70  microns long, i t s p o s t e r i o r t i p b e i n g rounded. are arranged protozoa  The f l a g e l l a  i n two s p i r a l s each p a s s i n g around the body of the  about three or four times.  The nucleus  d i s t a n t from the a n t e r i o r t a p e r i n g end.  i s two-fifths  -127Spironympha p o r t e r i . for  this- species or f o r S.  No  d e s c r i p t i o n was a v a i l a b l e  ovalis.  Micro,j oenia r a t c l i f f e i .  This i s a small p y r i f o r m -  shaped protozoa with the a n t e r i o r end f l a t t e n e d on one  side.  There are s i x t e e n to twenty f l a g e l l a . Torquenympha o c t o p l u s . microns.  This average about 11 by  There are eight ovoid parabasal bodies  20  i n the para-  basal ring* Trichonympha c o l l a r i s . i s the l a r g e s t species of t h i s genus v a r y i n g between 168 and parabasal  360 microns i n l e n g t h .  cords pass c l o s e to or touch the Trichonympha campanula.  long as the above species but may fact  that the parabasal  This species i s n e a r l y as be  separated  t h i r d s are s p h e r i c a l with  The parabasal  The p o s t e r i o r  two-  The parabasal  cords meeting j u s t behind  to 215 microns long.  This i s the smallest of  four species being 55--115 microns l o n g .  the  termitis.  p a r t s , the epimerite, protomerite i s a small c y l i n d r i c a l  nucleus.  cords are mostly f r e e  It i s 108  Trichonympha a g i l i 3 .  Hirmocystis  the  the a n t e r i o r p o r t i o n t a p e r i n g to a .;  from contact with the nucleus.  all  from i t by  This i s the smallest, species  of t h i s genus o c c u r r i n g i n Zootermopsis..  t a p e r i n g at each end.  nucleus.  cords u s u a l l y do not touch the  Trichonympha s p h a e r i c a .  s l i g h t l y rounded p o i n t .  A l l the  papilla.  the  I t i s spindle-shaped  ,  apparatus i s b a s k e t - l i k e ,  nucleus. T h i s gregarine c o n s i s t s of and deutomerite.  The  three  epimerite  -128KoFoidina ovata. The \)ody  i s rudimentary.  In t h i s gregarine is  the epimerite  non-septate.  D i s t r i b u t i o n w i t h i n the D i g e s t i v e T r a c t . With the exception of the gregarines a l l the  protozoa  ar-e l o c a t e d i n the hind i n t e s t i n e "between the small i n t e s t i n e and  the colon*  The  gregarines are always found  i n the mid i n -  t e s t i n e u s u a l l y s l i g h t l y p o s t e r i o r to the g a s t r i c caeca.  These  Sporozoa are e i t h e r attached to the "body w a l l or are f r e e swimming at which time the epimerite i s l a c k i n g *  Of the  protozoa  f e e d i n g i n the hind i n t e s t i n e the m a j o r i t y are i n the lumen, but a few  are attached to the w a l l of the i n t e s t i n e .  l i v i n g forms are:  a l l s p e c i e s of Trichonyrapha,  The f r e e  Torquenympha  o£tup_lus,- Micro j q enia r a t c l i f f e i , Trichomonas t ermopsidi s , Trieereomitus blomastix  termopsidis and Hexamastix t ermopsidis.  s t r i x i s usuallj^ found  attached to the w a l l of the  a n t e r i o r p o r t i o n of the hind gut but may the lumen.  Stre-  be f r e e l y swimming i n  Species of Pyrsonympha and Dinenympha f i m b r i a t a  appear f r e e both  i n the lumen and l i n i n g of the w a l l of the gut.  Cell Division. I have not observed in the process of d i v i d i n g . immediately  any of the species above l i s t e d However, such probably  occurs  f o l l o w i n g ecdysis when the fauna i s being r e s t o r e d  to i t s normal number. P h y s i o l o g i c a l R e l a t i o n s h i p between Protozoa and By examining the protozoa  Termite.  carefully, certain  species  -129are fcnind to contain f i b r e s and Such species are:  other small p i e c e s of wood.  a l l species of Trichonympha. Trichomonas  termopsidis, Spironympha p g r t e r i , Spirotrichonympha f l a g e l l a t a . Dinenympha f i m b r i a t a , Pyrsonympha ma,j or , T r i c e r c o m i t u s t ermops i d i s , Hexamastix termopsidis d i g e s t i n g wood.  l o s e of wood or paper(f25-) i oxygenation he was  cellu-  By a, combination of s t a r v i n g and  T h i s species i n the absence of the other  fla-  i n number r e s u l t i n g i n the death of the  This protozoan i s not a symbiont probably  nourishment from i t s host was  s u r v i v e on a diet of the  able to e l i m i n a t e s i l l the f l a g e l l a t e s but  g e l l a t e s diminished host.  are  Cleveland has)shown that when termites are  defaunated they were unable/to  Streblomastix.  These species d e f i n i t e l y  receiving i t s  or from the other protozoa  able to l i v e i n the presence of Trichomonas.  i n a c a r e f u l s e r i e s of experiments has  since i t  Hungate  ('38)  shown that Zootermopsis  can o b t a i n unaided^a maximum of approximately  one-third of the  t o t a l amount of m a t e r i a l that i s already s o l u b l e or that becomes s o l u b l e w i t h i n i t through d i g e s t i o n .  The  d i g e s t i o n of  two-  t h i r d s of t h i s t o t a l amount of m a t e r i a l i s dependent upon the protozoa. d i g e s t e d by  The t e r m i t e a l s o o x i d i z e s some of the m a t e r i a l the protozoa.  a c t i n g on c e l l u l o s e  Hungate a l s o showed that enzymes  (cellula'ses) are not  hind gut but are formed by the protozoa. are dependent on the food taken i n by a period previous  secreted by the termite Some of the  protozoa  the termite s i n c e , during  to e c d y s i s , the termite undergoes s t a r v a t i o n  with the r e s u l t a n t l o s s of at l e a s t a l l the l a r g e r species,  -130those of Trichonympha d i s a p p e a r i n g f i r s t .  The relationship..:  between the termite and the fauna of i t s hind i n t e s t i n e i s is- symbiotic i n the case of the m a j o r i t y of the s p e c i e s . The r e l a t i o n s h i p between the protozoa of the mid i n t e s t i n e and the termite i s a p a r a s i t e host  relationship.  However, these p a r a s i t e s have no apparent e f f e c t upon the c e l l s of  the i n t e s t i n e , nor i s the t e r m i t e v i s i b l y a f f e c t e d "by the  presence of l a r g e numbers of gregarines(Henry '33). Protozoan The  P o p u l a t i o n i n v a r i o u s Stages of the Host. t o t a l protozoan  p o p u l a t i o n o c c u r r i n g i n the hind  i n t e s t i n e v a r i e s with the i n s t a r , with the time since the l a s t  4',  moult and with the nearness to'''the next Newly hatched  moult.  nymphs do not have any protozoa.  In  Ret i c u l i t ermea hesperus they may a c q u i r e protozoa by proctodaeal feeding. first  In the case of gootermopsis the young nymphs of the  i n s t a r a r e f e d stomodaeally  by o l d e r nymphs or a d u l t s .  On the next moult the second i n s t a r nymph g r a d u a l l y r e c e i v e s more p r o c t o d a e a l food, e v e n t u a l l y r e c e i v i n g nothing e l s e . ing  Dur-  t h i s p e r i o d of p r o c t o d a e a l f e e d i n g the young nymph gradually  a c q u i r e s a protozoan  fauna.  A l l o l d e r nymphs and a l l castes of  both s p e c i e s have the same species of protozoa, l a t i o n probably  the t o t a l popu-  v a r y i n g c o n s i d e r a b l y s i n c e the d i g e s t i v e t r a c t  undergoes a c o n s i d e r a b l e i n c r e a s e i n s i z e .  The r e l a t i v e  propor-:  t i o n s of the v a r i o u s species of protozoa v a r i e s with the i n s t a r , old  nymphs having much l a r g e r numbers of species of Trichonympha  than second or t h i r d i n s t a r nymphs.  -131Thr'ee t o f o u r weeks b e f o r e a period'of self-starvation, i n t e s t i n e disappears  s t a r v e and d i s a p p e a r . first  so t h a t t h e f o o d  but the protozoa  cht  t h e i r food supply  remain.  o f f , the wood-digesting Of t h e s e ,  content  enters  of the  However, w i t h protozoa  gradually  t h e s p e c i e s o f T r i c h o n y m p h a go  f o l l o w e d by T r i c h o m o n a s i n t h e c a s e o f Z o o t e n u o p s i s _ .  that a l l of these a r e absent before other  ecdysis the termite  t h e moult o c c u r s .  so  Of t h e  s p e c i e s , S t r e b I o m a s t i x a n d H e x a m a s t i x may^ o r may n o t r e -  main t h r o u g h  the p e r i o d of the moult and T r i c e r c o r a i t u s always  p e r s i s t s vl-n-^t-he—i-ntestf-ifl-e'. Following ecdysis thepopulation i s gradually restored to i t s o r i g i n a l p r o p o r t i o n s . Alates previous and  to moulting  l o s e most o f t h e i r  acquire the f u l l population only after  protozoa  swarming.  C a r e f u l s t u d i e s o f t h e number o f p r o t o z o a  i na large  n o r m a l nymph have shown t h a t t h e h i n d i n t e s t i n e may c o n t a i n as many a s 25,000 T r i c h o n y m p h a a n d a b o u t 500,000 T r i c h o m o n a s (Andrew *30). Factors A f f e c t i n g Protozoan  Content.  The c h i e f f a c t o r r e s p o n s i b l e f o r a p o p u l a t i o n o f protozoa  i s the presence of food w i t h i n the d i g e s t i v e t r a c t .  As was s t a t e d a b o v e ^ s t a r v a t i o n o f t h e h o s t with the s t a r v a t i o n of the protozoa. may b e s u c h t h a t a t l e a s t Should and  reason  related  The d e g r e e o f s t a r v a t i o n  the l a r g e r species  f u n g i be a p a r t i a l  there i sl i t t l e  i sclosely  source  t o doubt t h i s ,  disappear.  of food f o r protozoa, s t a r v a t i o n o f the host  -132would r e s u l t - a s b e f o r e  i n t h e l o s s o f many o f t h e f l a g e l l a t e s .  . ..Experimentally by  oxygenation  t o s u c h an e x t e n t  content  may "be a l t e r e d  that the d i g e s t i v e tract i s  f r e e o f protozoaiCC^f^e^^d^J-p j  entirely  The effect  the protozoan  a c t u a l process  of moulting  the protozoan content.  or ecdysis  do/es n o t  I t i s the conditions  attendant  upon e c d y s i s w h i c h a r e r e s p o n s i b l e f o r t h e change i n p o p u l a t i o n , that  is_host starvation. ReFaunat i o n f o l l o w i n g E c d y s i s * Previous  to the time o f e c d y s i s  i s reduced to a very  small part  content  o f i t s f o r m e r p o p u l a t i o n so  that refaunation or the a c q u i s i t i o n outside  the protozoan  of protozoa  from  sources  t h e i n d i v i d u a l must t a k e p l a c e f o r t h e c o n t i n u e d  ment o f t h e t e r m i t e .  A s was s t a t e d above t h e r e may s t i l l  three  i n the hind  The  species present  intestine during  a d d i t i o n s t h e r e f o r e consist of the other  method o f r e f a u n a t i o n  i s not d e f i n i t e l y  That  does n o t t a k e p l a c e by a n a l c o n t a c t w i t h p r o t o z o a shown by Andrew ( ' 3 0 ) .  Refaunation  definitely  can b e n o r m a l l y  is and  i n motile  some i n d i c a t i o n  obtained  or feces i s  f o r t h i s f e e d i n g were n o t F a e c e s do n o t  c o n d i t i o n o r i n c y s t s , though  t h a t c y s t o i d forms of Trichomonas  Trichonympha campanula e x i s t  unfavorable  conditions.  refaunation  The methods by w h i c h  d e t e r m i n e d i n Andrew's e x p e r i m e n t s .  contain protozoa  The e x a c t  d o e s t a k e p l a c e r e a d i l y by  f e e d i n g on m a t e r i a l c o n t a i n i n g p r o t o z o a * the p r o t o z o a  be  ecdysis.  species.  known.  develop-  there  termopsidis  i n t h e i n t e s t i n e under c e r t a i n  F e e d i n g t e r m i t e s on f a e c e s  does n o t  r e f a u n a t e them. but  C a n n i b a l i s m l e a d s to a t r a n s f e r of p r o t o z o a  i t i s ;not t h e o n l y a g e n c y t h a t a c h i e v e s t h i s r e s u l t .  tozoa haveibeen  Pro-  f o u n d i n some i n s t a n c e s on t h e u n e x t r u d e d  and i n t h e f i r s t  drop o f i n t e s t i n a l f l u i d ,  pellet  hence t h e t r a n s f e r  o f p r o t o z o a d u r i n g p r o c t o d e a l f e e d i n g may b e one o f t h e n o r m a l methods o f r e f a u n a t i o n .  Andrew s t a t e s t h a t  i n mixed  groups of  f a u n a t e s and d e f a u n a t e s w h e r e c a n n i b a l i s m was p r e v e n t e d and no stomodeal  f e e d i n g o r d r o p p i n g o f l i q u i d s was o b s e r v e d , p r o c t o d e a l  f e e d i n g on t h e p a r t o f b o t h f a u n a t e s and d e f a u n a t e s  continued  as u s u a l , b u t r e f a u n a t i o n - w a s not a c c o m p l i s h e d , i n d i c a t i n g  that  t h i s may  In a  few  n o t be t h e most e f f e c t i v e method o f r e f a u n a t i o n .  c a s e s , l i q u i d d r o p p i n g s were found to c o n t a i n l i v i n g  lates, which  flagel-  so t h a t f e e d i n g on t h e s e and on f r e s h n e s t s t r u c t u r e s i n intestinal  fluid  i n p a r t f o r normal  i s u s e d a s a cement may p r o b a b l y  r e f a u n a t i o n s . B y , u s i n g r e c e n t l y moulted i n -  d i v i d u a l s i t Yvas d e t e r m i n e d a t i o n has u s u a l l y Effect  account  that  i n the normal  colony, refaun-  t a k e n p l a c e by the t h i r d o r f o u r t h day.  on P r o t o z o a c a u s e d b y f e e d i n g T e r m i t e s on Woods  O t h e r t h a n t h o s e p r e f e r r e d by - them.. In a s e r i e s of experiments used p r e f e r e n c e of t e r m i t e s , heartwood  to d e t e r m i n e  t h e wood  c e d a r was one o f t h e woods u s e d .  I t was f o u n d t h a t when nymphs o f Z o o t e r r n o p s i s a n g u s t i c o l l i s w e r e g i v e n c e d a r and some o t h e r wood, neglected, only the s l i g h t e s t a b l e on i t . attempt  t h e c e d a r was  t r a c e of tunneling being notice-  The t e r m i t e a p p a r e n t l y w o u l d  to eat cedar.  invariably  As a r e s u l t ,  r a t h e r s t a r v e than  s t a r v a t i o n and t h e d i s a p p e a r a n c e  -134o f many o f t h e p r o t o z o a P a r a s i t i c Protozoa  will  be i n e v i t a b l e .  a s a Means o f C o n t r o l .  Of a l l t h e f o r m s e x i s t i n g i n t h e t e r m i t e canal  only  t h e g r e g a r i n e s j_j___j___the power t o c o n t r o l  H o w e v e r , t h e s e p a r a s i t e s h a v e no a p p a r e n t  gregarines.  termites.  e f f e c t upon t h e c e l l  of t h e i n t e s t i n e , n o r i s the t e r m i t e v i s i b l y p r e s e n c e o f l a r g e numbers o f  alimentary  a f f e c t e d by t h e  PART IV FlIiTGI AND THEIR RELATION TO The  TERMITES  Fungi* F u n g i a r e a l w a y s f o u n d i n c o l o n i e s o f Zoot ermopsis  and  often  i n c o l o n i e s o f RjgtJ^cuI i t erme_s *  Of t h e s e , some a r e  wood<=-destroying h u t a l a r g e number a r e s e c o n d a r y f u n g i ing  o r l i v i n g on wood a l r e a d y  isolated  some t w e n t y - f i v e  decaj^ed.  Hendee ('33)  has  genera from c o l o n i e s of R e t i c u l i t e r m e s  h e s p e r u s and twenty-two from Zooterrnopsis t h i s connection  attack-  angusticoilis.  none w e r e f o u n d s p e c i f i c f o r a d e f i n i t e  In species  o f t e r m i t e , h u t a l a r g e number were f o u n d common t o t h e c o l o n i e s of s e v e r a l  species* These f u n g i were m o s t l y those b e l o n g i n g  Imp e r f e c t i . ray c e t e s  to the Fungi  S m a l l numbers o f them w e r e PhycomVy c e t es,  or Basidiomycetes.  In t h i s l a s t  Asco-  named group o c c u r t h e  wood^-destroying f u n g i . U s u a l l y Zootermopjais i s seldom found a s s o c i a t e d l i v i n g B a s i d i o m y c e t e s b u t i s o f t e n i n wood d e s t r o y e d wood-consuming f u n g i and i n h a b i t e d by secondary Within tunnels,  the colony  extending a short  distance frasa,  into  the surface  T e r m i t e s on e n t e r i n g a p i e c e  present  saprophytes.  t h e wood.  o f t h e t e r m i t e s a n d even f r o m t h e i r d i g e s t i v e  those f u n g i already  by t h e  t h e f u n g i a r e f o u n d a l l a.long t h e  may be i s o l a t e d f r o m p e l l e t s ,  or hyphae from t h e o u t s i d e  with  of the bodies  tracts.  o f wood may  to the inner p a r t s . i n t h e wood.  The f u n g i  carry  spores  They may  spread  -136Fungi their  a r e b y no means d e p e n d e n t on t e r m i t e s f o r  spread but they a r e a i d e d  tunnels provide a moist  c o n s i d e r a b l y by them.  The  atmosphere r i c h i n carbon d i o x i d e  f a v o r a b l e to the growth of f u n g i . Fungi  as a Source of Food. I f t e r m i t e s a r e f e d on f u n g u s - c o n t a i n i n g wood  there  i s an a p p r e c i a b l e i n c r e a s e i n w e i g h t a n d i n t h e amount o f nitrogen present  i n t h e body.  I f t h e y a r e f e d on  wood t h e r e i s u s u a l l y no I n c r e a s e is prevalent.  Should  tic  i n weight but h i g h m o r t a l i t y  t h e r e be an I n c r e a s e  i n c r e a s e i n n i t r o g e n content  fungus-free  i n w e i g h t and an  i t i s due t o e x c e s s i v e c a n n i b a l i s  feeding* An i n c r e a s e i n a v e r a g e w e i g h t o f t e r m i t e s may  f r o m f e e d i n g them on f u n g u s - f r e e  result  wood on w h i c h t h e r e i s a  s u p e r f i c i a l g r o w t h o f some f u n g u s .  However Z o o t e r m o p s i s  exist  •A  b e t t e r on a d i e t chemical  o f r o t t e n wood.  I t seems p r o b a b l e  that the  c h a n g e s b r o u g h t a b o u t by t h e f u n g i a r e i m p o r t a n t i n  p r e p a r i n g t h e wood t o be a s u i t a b l e d i e t f o r Z o o t e r m o p s i s . Termites  a r e u n a b l e t o l i v e a n d grow n o r m a l l y  diet of p u r i f i e d c e l l u l o s e  (Cook a n d S c o t t  '33).  on a  PART V ECONOMIC STATUS Type o f Work and E x t e n t  o f Damage.  S i n c e R e t i c u l i t e r m e s h e s p e r u s has up  to the  present  C o l u m b i a <m±y- "been re,c_o.rded d o i n g d a m a g e ^ o  time  in British  root  c e l l a r ^ the remarks to f o l l o w concern  p o s s i b i l i t i e s f o r damage by s o u t h e r n Okanagan w h e r e t h e y  Zoot ermopsis.  the small species are f a i r  a  The i n the  a r e f a v o u r e d by h i g h a v e r a g e  temperatures. The ted  work o f ' Z o o t e r m o p s i s  i n this province i s restric-  to the s t r u c t u r e s a l r e a d y s t a r t e d  are u n i f o r m i l y quite moist.  t o d e c a y and  those  Such s t r u c t u r e s t h a t have  w i t h t h e g r o u n d a r e t h o s e most f a v o u r e d by  contact  this termite.  t h e s e , wooden f o u n d a t i o n s , p o l e s and p o s t s o f a l l k i n d s t i e s s u f f e r most f r o m t h e i r a t t a c k s .  which  Of and  Cement f o u n d a t i o n s f o r  wood s t r u c t u r e s g i v e b e t t e r i n s u r a n c e a g a i n s t a t t a c k s , b u t t h e y must be  o f s u c h a s i z e and  t h e wood i t s e l f on  h e i g h t above the ground that  d o e s n o t f r e e l y p i c k up m o i s t u r e .  cedar p o l e s a r e c o n f i n e d c h i e f l y  ground l e v e l .  C o l o n i e s h o w e v e r may  The  attacks  t o t h e sapwood b e l o w extend  at l e a s t  the  f i v e feet  above g r o u n d on t h e n o r t h e r n e x p o s u r e .  T i e s made o f  h e m l o c k o r D o u g l a s f i r i f not  i n v a r i a b l y become d e s -  t r o y e d by  creosoted  either  t h e dampwood t e r m i t e * The  e x t e n t o f damage by  Zootermopsis  in this  i s b e i n g f e l t more and more, p r o b a b l y  through  more c o o p e r a t i o n w i t h e n t o m o l o g i s t s .  That t h e damage i s i n -  creasing i s questionable.  education  province and  Damage t o d w e l l i n g s i s c o n f i n e d t o  -138t h o s e b u i l t many y e a r s ago when cement f o u n d a t i o n s w e r e s e l d o m used. Y/ood P r e f e r e n c e s . T h e r e a r e two methods by w h i c h wood p r e f e r e n c e o f t e r m i t e s can be determined.  One o f them i s t h e s e t t i n g out  o f u n i f o r m - s i z e d b l o c k s i n s u i t a b l e l o c a t i o n s and t h e n a w a i t i n * "te?rmiTe-'3>  t h e a t t a c k by them.  T h i s method has b e e n u s e d where t e r m i t e s  form an a p p r e c i a b l e p e r c e n t a g e of i n s e c t p o p u l a t i o n - t h e t r o pics*  The o t h e r method w h i l e b e i n g p u r e l y a r t i f i c i a l  most f e a s i b l e one f o r t h i s  i s the  province.  In the experiment conducted, a s e r i e s of b l o c k s of green cedar, that  f i r and hemlock were used*  each p a i r  c o n s i s t e d o f two s p e c i e s .  These w e r e p a i r e d so I n t h e s i d e of each,  a h o l e was b o r e d so t h a t when t h e i r f a c e s were brought t o gether  a cylindrical  c a v i t y was f o r m e d .  the purpose of a t u n n e l . b l o c k s were p l a c e d sis angusticollis. Having thus  This cavity, served  Into the c a v i t y of each p a i r of  t e n nymphs o f t h e s e v e n t h  instar  E a c h e x p e r i m e n t was done i n t r i p l i c a t e .  introduced the t e r m i t e s into the c a v i t y  b l o c k s were t i g h t l y  Zooterrnop-  t h e two  f a s t e n e d t o g e t h e r w i t h s t r i n g so t h a t t h e  nymphs c o u l d not e s c a p e .  A l l o f t h e s e b l o c k s were then  placed  i n two l i k e c o n t a i n e r s so t h a t t h e e x t e r n a l c o n d i t i o n s were t h e same f o r a l l .  Observations  w e r e t h e n made o v e r a p e r i o d  o f a b o u t f i v e o r s i x w e e k s , each t i m e n o t i n g t h e t o t a l present  number  a n d t h e number d y i n g o r d e a d , and t h e amount o f t u n -  n e l i n g made i n each b l o c k o f t h e p a i r .  These o b s e r v a t i o n s a r e  -139recorded  i n the appendix. While the c o n d i t i o n s i n these experimental  "blocks  a r e f a r f r o m i d e a l and w h i l e no wood d e s t r o y i n g f u n g i were t ^— present  t h e e x p e r i m e n t s were c o m p a r a b l e .  P e n i c i l l i u m was t h e  fungus which developed w i t h i n t h e b l o c k s . The c o n c l u s i o n s  from t h i s s e r i e s of experiments are  as f o l l o w s : 1. D o u g l a s f i r i s p r e f e r r e d  to e i t h e r cedar or  h e m l o c k , and h e m l o c k i s p r e f e r r e d t o c e d a r , as shown by t h e s i z e and l e n g t h o f t u n n e l s . 2. I n wood w h i c h i s f e d upon r e l u c t a n t l y , t h e termites  i n v a r i a b l y become weakened  nibalism  resulting.  with.can-  I n t h i s e x p e r i m e n t w h i c h was t o h a v e b e e n t h e b a s i s for  others,  i t was r e a l i z e d t h a t t h e l e n g t h o f runways o r  t u n n e l s and, t h e s u r v i v a l o f t h e t e r m i t e s a r e a f f e c t e d n o t o n l y by  the species  o f wood t e s t e d b u t a l s o by t h e s p e c i f i c g r a v i t y ,  r a t e of growth, hardness, contents  a f f e c t i n g t a s t e and  v a l u e , and t h e age o f t h e wood w i t h i n a  food  species.  CONTROL Zoot e r m o p s i a a n g u s t i c o l i i s and n e v a d e n s i s may c o n t r o l l e d by t h e u s e o f cement f o u n d a t i o n s  and t h e  be  chemical  t r e a t m e n t , o f t h o s e s t r u c t u r e s w h i c h must come i n c o n t a c t the s o i l . pouring  They may be k i l l e d  poison  dusts  with  i n c o l o n i e s most e f f e c t i v e l y by  s u c h as s o d i u m f l u o r i d e o r s o d i u m  silico  f l u o r i d e i n t o the t u n n e l s ^ f o l l o w i n g which the opening i s  -140ealed.  I n . t h i s way  t h e i r h a b i t s of grooming are made use  of.  -1BIBLIOGRAPHY 1.  Andrew, B . J . 1930.  Method and r a t e of protozoa r e f a u n a t i o n i n the termite, Termopsis a n g u s t i c o i l i s . Univ. C a l i f . P u h l . Z o o l . , 33: pp 449-470  2.  Balfour-Browne, P. 1932.  3.  A Textbook of P r a c t i c a l Entomology.  Banks, ITathan. 1920.  A R e v i s i o n of the H e a r o t i c Termites. 108.  4.  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T e r m i t e s and T e r m i t e C o n t r o l .  Univ.  Calif.  Press. 33.  K u d o , R. R. 1931,  34".  of  Protozoology.  L e e c h , Hugh B. 1938.  35.  Handbook  L e t t e r dated March, 1 4 t h .  L i g h t , S. F,, and S a n f o r d , 1928,  Experimental  M.  F.  transfaunation  C a l i f . P u b l . Z o o l . , 3 1 : pp 36.  of t e r m i t e s . Univ. 269-274.  L u n d , E. 1930.  The e f f e c t  o f d i e t upon t h e i n t e s t i n a l  Termopsis.  fauna of  U n i v . C a l i f . P u b l . Z o o l . , 36: pp  81-96. 37.  Mac G i l l i v r a y , A l e x . 1923.  38.  39.  External Insect  M i a l l , L . C., 1886.  P.  and Denny,  The  Anatomy.  D.  Cockroach.  IToyes, B. 1930.  The P e r i p h e r a l S e n s e O r g a n s i n t h e Termopsis a n g u s t i c o l l i s P u b l . Z o o l . , 3 3 : pp  40.  P o w e l l , W. 1928.  (Hagen).  Termite,  Univ.  Calif.  259-286.  H. On t h e m o r p h o l o g y o f P y r s o n y m p h a w i t h a d e s c r i p t i o n o f t h r e e new hesperus (Banks). 31: pp  179-200.  species from R e t i c u l i t e r m e s  Univ. C a l i f . Publ.  Zool.,  41.  Report  f o r 1936.  42.  Snodgrass, 1909.  Climate of B r i t i s h  Columbia.  R. E. The T h o r a x the Wings.  o f I n s e c t s and t h e A r t i c u l a t i o n o f P r o c . U.S. N a t . Museum, 36: pp 5 1 1 -  595. 43'.  1927. Morphology  and Mechanism o f t h e I n s e c t  Thorax.  S m i t h s . M i s c . C o l l . , 80: N o . 1. 44.  1930.  How I n s e c t s E l y . S m i t h s R e p o r t f o r 1 9 2 9 . pp 3 8 3 - 4 2 1 .  45. 46.  1935. P r i n c i p l e s o f I n s e c t Snyder,  Morphology,  T. E.  1924.  "Adaptations" to Social L i f e : (Isoptera) Smiths. Misc. C o l l . ,  The T e r m i t e s 76: No. 1 2 ,  pp 1-14. 47. 48*  1935.  S t r i c k l a n d , E. H. 1938,  49.  Our Enemy t h e T e r m i t e ,  L e t t e r dated March 18th.  Sumner, E t h e l 1933.  Craig*  The S p e c i e s o f t h e T e r m i t e Genus E m e r s o n (n T e r m o p s i s  Hagen).  Zootermopsis  Univ. C a l i f . Publ.  E n t . , 6: No. 7, pp 197-230, 50.  Swezy, O. 1919.  The p s e u d o p o d i a l method o f f e e d i n g by T r i c h o nymphid f l a g e l l a t e s p a r a s i t i c termites. 391-400.  i n wood-eating  U n i v . C a l i f . P u b l . Z o o l . , 20: pp  -750A.  Thompson, C. B. 1913.  Study  o f t h e B r a i n o f Three Genera o f A n t s ,  w i t h S p e c i a l R e f e r e n c e t o t h e Mushroom B o d i e s Jo urn. 51.  Comp. Z.00I., 2 3 .  1 9 1 6 . The B r a i n a n d F r o n t a l G l a n d o f t h e C a s t e s o f t h e 'White A n t ' , L e u c o t e r m e s Jo u r n .  52.  flavipes,  Kollar.  o f Comp. N e u r . , 2 6 : pp 553-603.  1 9 1 8 . The O r i g i n o f t h e C a s t e s o f t h e Common T e r m i t e , Leucotermes  flavipes Kol.  J o u r n . o f Morph,,  30: pp 83-134. 53.  1 9 1 9 . The D e v e l o p m e n t o f t h e C a s t e s o f M i n e Genera and T h i r t e e n S p e c i e s o f T e r m i t e s .  Biol.  Bull.,  36: pp 379-398. 54.  Thompson, G. B., a n d S n y d e r , 1920.  T. E.  The ' T h i r d Form', t h e W i n g l e s s R e p r o d u c t i v e . Type o f T e r m i t e s : hinotermes.  55.  R e t i c u i i t e r m e s and Phor-  J o u r n . o f M o r p h . , 3 4 : pp 591-633.  1 9 2 2 . The C a s t e s o f T e r m o p s i s .  J o u r n . o f Morph.,  36: pp 4 9 5 - 5 3 6 . 56.  U v a r o v , B. P. 1928.  57.  Walker,  I n s e c t N u t r i t i o n and Metabolism. of  the Literature.  pp  255-343.  A Summary  T r a n s . E n t . Soc*  London,  E. M.  1919.  The T e r m i n a l A b d o m i n a l teroid Insects:  S t r u c t u r e s o f Orthop-  A P h y l o g e n e t i c Study.  E n t . S o c . o f Am., 1 2 : No. 4, pp 267-316.  Annals,  58.  1931.  On  the Clypeus  and L a b i u m o f P r i m i t i v e I s e c t s .  C a n a d i a n E n t o m o l o g i s t , 63: pp 59.  Weber, Hermann. 1933.  Lehrbuch  der Entomologie.  F i s c h e r i n Jena. 60'*  61. 62.  W h e e l e r , W.  V e r l a g von  Gust  ppl26-127.  M.  1928.  The  1937.  Mosaics  Wigglesworth, 1934.  63.  75-81.  V.  Social Insects. and  other Anomalies  amon| A n t s .  B.  Insect Physiology.  Yuasa, Hachiro. 1920.  The  Anatomy o f t h e Head'and M o u t h P a r t s o f  O r t h o p t e r a and E u p l e x o p t e r a . 33: Ho.  2, pp  251-307.  J o u r n . o f Morph.,  -141C01TCLUSI0F. , ,In c o n c l u d i n g  this thesis a l i s t  of the problems  w h i c h I e n c o u n t e r e d a n d d i d n o t have t i m e t o c o m p l e t e w i l l "be given  here.  They a r e a s f o l l o w s :  A s s y m e t r y i n t h e e p i p h a r y n x and h y p o p h a r y n x . Function Histology  of the f i r s t  abdominal s p i r a c l e s .  of the alimentary of  c a n a l b e f o r e and a t t i m e  ecdysis.  Histology  o f t h e m a n d i b u l a r and m a x i l l a r y  Histology  of the b r a i n of Zootermopsis.  Histological Histology  the  study o f t h e development o f t h e b r a i n .  o f the r e p r o d u c t i v e  especially  glands.  systems o f  Zooterrnopsis.  t h o s e o f t h e male i n o r d e r t o d e t e r m i n e  exact i d e n t i t y o f each p a r t .  Histology  of the gland  located  on a l l t h e t i b i a e .  The  s t r u c t u r e and f u n c t i o n  of the corpora a l a t a .  The  complete study o f t h e embryology o f  a n g u s t i c o l l i s to determine i f p o s s i b l e  Zooterrnopsis the source  of d i f f e r e n t i a t i o n o f t h e c a s t e s . In t h i s t h e s i s considerable on  emphasis has been  placed  t h e e x t e r n a l m o r p h o l o g y o f Zo otermop s i s angu s 11c o 1 1 i s and  Z. n e v a d e n s i s t o f i n d species.  the d i f f e r e n c e s  e x i s t i n g , b e t w e e n t h e two  The m o r p h o l o g y o f t h e v a r i o u s  well investigated.  In addition  castes  has been  fairly  to the other d e s c r i p t i o n s , i n -  t e r n a l anatomy a n d i n t e s t i n a l f a u n a , a b e g i n n i n g has been made in  t h e embryo l o g i c a l  s t u d y o f Zoo t e r m o p s i s an gu s t i c o l _ l i s .  -142-  Append WOOD PREFERENCE TESTS B l o c k s o f D o u g l a s F i r , H e m l o c k and c e d a r were o b t a i n e d i n t h e g r e e n a n d w e r e e x p o s e d t o t h e a t m o s p h e r e f o r 14 days so t h a t t h e p i e c e s w o u l d be i n t h e same c o n d i t i o n o f ' m o i s t u r e c o n t e n t . The p i e c e s were a p p r o x i m a t e l y 3 X 5 X 1" i n d i m e n s i o n each w i t h a h o l e 3/4" i n d i a m . b o r e d t o a d e p t h o f 3/8 - l / 2 " . The b l o c k s were then a r r a n g e d i n p a i r s , t h e h o l e s c o i n c i d i n g to form c y l i n d r i c a l chambers i n t o w h i c h t e n s e v e n t h i n s t a r nymphs o f Zoot errnop s i s a n g u s t i c o l i i s were p l a c e d . The p a i r e d b l o c k s were t h e n t i e d t o g e t h e r and p l a c e d i n b a t t e r y j a r s w h i c h were almost a i r - t i g h t . T h r e e l o t s o f each s e t o f b l o c k s w e r e u s e d to r e d u c e t h e p o s s i b i l i t y of. e r r o r . A n o t h e r s e t o f b l o c k s was p l a c e d i n a t e r m i t e c o l o n y c o n t a i n i n g p i e c e s o f d e c a y e d wood. The e x p e r i m e n t was s e t up w e r e a r r a n g e d as f o l l o w s : 3. c e d a r 1. c e d a r 2. f i r 4, f i r  7. c e d a r 8. hemlo ck is;.' . f i r 14. h e m l o c k 19. c e d a r  9. 10. 15. 16. 20.  30.XI.1936 a n d 5. 6.  c edar f ir  cedar hemlo ck  11. c e d a r 12. h e m l o c k  fir hemlo ck  17. 18. 21.  fir  the b l o c k s  fir hemlock hemlock (used i n c o l o n y )  -143-  ;  No. o f B l o c k  Observations Ho. i n e a c h Block  on -wood p r e f e r e n c e Total  tests Remarks  1. 2.  0 9-1  3. 4.  0 10  10  5. 6.  Q 10  10  N a r r o w t u n n e l a r o u n d r i m i n No.6  7. 8.  1 9  10  T u n n e l 6 mm.  9. . • 10.  0 10  10  11. 12.  0 9  9  One t e r m i t e p a r t l y e a t e n , abdomen l a c k i n g . T u n n e l i n No.12, 14 mm. long.  13. 14.  8 1  9  No t r a c e o f t h e t e n t h t e r m i t e . T u n n e l i n No. 16, 13mm. l o n g .  6 4  10  T u n n e l i n No. 16, 13mm.' l o n g .  9 1  10  Three t u n n e l s , l a r g e s t  15. 16. 17. 18.  .  9 One t e r m i t e d y i n g , p a r t l y eat en p a r t o f abdomen and l e g s m i s s i n g on one s i d e . Short  t u n n e l i n Mo. 4  l o n g i n No. 8.  14mm.  A l l p i e c e s s h o w i n g s i g n s o f m o l d on o u t s i d e o f t h e blocks. P i e c e s o p p o s i t e t h o s e h a v i n g t i x n n e l s show t h e o u t l i n e of the t u n n e l s . B l o c k s p l a c e d i n c o l o n y cemented t o g e t h e r w i t h d e c a y i n g wood o f t h e c o l o n y . Observation  made on 8 . X I I I . 1 9 3 6 .  -144-  . , ITo. o f B l o c k  O b s e r v a t i o n s on -wood p r e f e r e n c e t e s t s Ho. I n e a c h Block  Total  Remarks  1. 2.  0 9  9  B o r i n g i n t h e sunmerwood i n Ho. 2.  3. 4.  1 8  9  Head a n d t h o r a x o f 1 0 t h e a t e n . T u n n e l i n Ho. 4, 5ram. Two o t h e r s s t a r t e d i n t h e spring-wood.  5. 6.  0 10  10  T u n n e l i n Ho. 6, 3mm. ~w«~e%he3?e Two o t h e r s s t a r t e d i n t h e s p r i n g wood, 1 0 t h p a r t l y e a t e n (abdomen). T u n n e l i n ITo. 8, 14mia.  7* 8,  .  '  1 8  9  9. 10.  0 10  10  11. 12.  0 9  9  T u n n e l i n ITo. 1 2 , 28mm. i n t h e s p r i n g - w o o d . 2nd t u n n e l 4mm.  13. 14.  8 1  9  T^o t u n n e l s i n No. 14, 6 a n d 9 mm.  15. 16.  7 3  10  T u n n e l s i n No. 1 6 , 5,6, a n d 22mm.  17. 18.  10 0  10  Two t u n n e l s s t a r t e d i n s p r i n g wood i n Ho. 1 7 . T u n n e l s i n No. 18, 3, 4, a n d 8mm.  T u n n e l i n Ho. 1 0 , 16mm.  T e r m i t e s i n b l o c k s 1 9 , 20 a n d 21 i n good  condition*  O b s e r v a t i o n made on 15.XII.1936.  Gbservations Ho.  of Block . ,, 1. 2.  -145on wood p r e f e r e n c e  tests  Ho. i n e a c h Block 0 8  Total 8  T u n n e l i n (2)  3. 4.  0 8  8  Remainder o f 9 t h . Tunnel i n (4) a c r o s s g r a i n .  5. 6.  0 10  7. 8.  0 6  6  Three d i e d , caiise u n c e r t a i n b u t p r o b a b l y due t o t o o m o i s t a c o n d i t i o n . Length o f tunnel i n ( 8 ) 11mm.  o 7  7  Three d i e d d i s c o l o r e d as i n 7 a n d 8. L e n g t h o f t u n n e l i n ( l O ) 16mm. I m p r e s s i o n o f t h i s tunnel i n (9).  0 8  8  9 t h s t i l l a l i v e ( a l t h o u g h headl e s s ) . Length o f tunnel i n (12) 38mm. I m p r e s s i o n o f t h i s t u n n e l i n ( l l ) . 2 n d t u n n e l i n (12) 4mm.  13. 14.  7 2  9  15. 16.  5 5  10  T u n n e l s i n (14) 20 a n d 6mm. T u n n e l i n (13) 5mm. I m p r e s s i o n s of these t u n n e l s i n o p p o s i t e block. T u n n e l i n (16) 23mm., 8mm. w i d e f o r i8mm. o f i t s l e n g t h . O t h e r t u n n e l s i n (16) 5 a n d 6mm. Impressions o f these i n (15).  17* 18.  8 2  10  9. 10.  11. 12,,  '  "  10  Remarks i n springwood.  Tunnel i n (6) i d e n t i c a l that of four.  T u n n e l s i n (18) T u n n e l s i n (17) springwood.  with  9,3, a n d 4mm. 3~5mm. deep i n  I t i s a p p a r e n t t h a t t h e c o m b i n a t i o n o f any wood w i t h cedar i s n o t f a v o r a b l e t o t h e h e a l t h o f t h e c o l o n i e s . I n b l o c k s 7ffi8 a n d 9 & 10 a number o f e a c h have d i e d p r o b a b l y n o t as a r e s u l t of c a n n i b a l i s m b u t t o o h i g h m o i s t u r e c o n d i t i o n s r e s u l t i n g i n f u n g u s attack. T h e s e s p e c i m e n s s h o w e d ' d i s c o l o r a t i o n o f the w i n g s , head and l e g s , t u r n i n g b l a c k . C e d a r h a s o n l y s l i g h t l y b e e n b o r e d a n d t h e n o n l y where i t was i n c o n t a c t w i t h t h e t u n n e l s i n t h e o p p o s i t e b l o c k . Observation  made on 21.XII.1936*  -146Observations o f Blo'tak  on wood p r e f e r e n c e  Ho. I n e a c h Block  Total  tests Remarks  1 2  0 6  6  Two p a r t l y e a t e n t e r m i t e s . Two t u n n e l s deepened i n ( 2 ) i n s p r i n g wood, d e p t h 9.5.  3 4  0 8  8  Four shallow tunnels i n s p r i n g wood, a n o t h e r a c r o s s g r a i n i n (4), l e n g t h 7.  5 6  0 8  8  Two s h a l l o w t u n n e l s i n s p r i n g wood, one a c r o s s g r a i n i n ( 6 ) , l e n g t h 5mm, ITo e v i d e n c e o f 9 t h and 1 0 t h t e r m i t e s .  7 8  0 5  5  6 t h dead as a r e s u l t o f f u n g u s a t t a c k ? Fungus g r o w i n g i n c e l l and t u n n e l . T u n n e l i n ( 8 ) 13mm.  9 10  0 5  5  Two dead p r o b a b l y a t t a c k e d b y f u n gus. L e n g t h of t u n n e l I n (10) 18mm. I m p r e s s i o n o f t h i s t u n n e l in (9).  11 12  0 5  5  Remnants o f t h r e e t e r m i t e s a l l turned black. Length of tunnel i n ( 1 2 ) 40mm. S e c o n d t u n n e l u n changed . F u n g u s g r o w t h i n c e l l .  13 14  7 2  9  T u n n e l s i n ( 1 4 ) 2 0 , 6 a n d 8mm. T u n n e l i n ( 1 3 ) 9n.:i.  15 16  3 7  10  17 18  6 3  9  Observation  T u n n e l s i n ( 1 6 ) 2 5 , (8.5mm. w i d e ) , o t h e r s unchanged. T u n n e l s I n ( 1 7 ) and ( 1 8 ) unchanged.  made on 2 8 . X I I . 1 9 3 6 .  -147Dbservations No.  of Block  Ho. i n e a c h Block  on wood p r e f e r e n c e Total  tests Remarks  1 2  0 6  6  One t u n n e l i n ( 2 ) h a s a l m o s t reached t h e outside. Three o t h e r s s t i l l shallow.  3 4  0 7  7  8 t h i n p a r t . Tunnels i n (4) l i t t l e changed.  5 6  0 7  7  Tunnels i n (6) l i t t l e 8th i n part.  7 8  0 5  5  T u n n e l u n c h a n g e d e x c e p t more f u n g u s g r o w t h i n c e l l a n d t u n n e l . One t e r m i t e weak.  9 10  0 4  4  5 t h d e a d , 4 t h weak. No change i n tunnels.  11 12  0 4  4  5 t h d i s c o l o r e d and p a r t l y eaten. 4 t h w i t h o n l y t h r e e l e g s and somewhat weak. T u n n e l s unchanged.  13 14  8 0  8  9 t h p a r t l y eat en. T u n n e l i n ( 1 3 ) , 10mm. a n d d e e p e r . T u n n e l s i n ( 1 4 ) , 6, 1 1 , & 21mm.  15 16  4 6  10  17 18  6 3  9  changed.  T u n n e l i n ( 1 6 ) a t end i s 7mm o t h e r s unchanged*  deep,  Unchanged.  D e s c r i p t i o n o f t e r m i t e dead i n (9-10). Head a n d i t s a p p e n d a g e s a l m o s t b l a c k i n c o l o r * Wings d a r k brown. L e g s brown_ to b l a c k i n c o l o r . T h o r a c i c segments b e c o m i n g d a r k . Abdomen s t i l l of n a t u r a l c o l o r . T h i s d e s c r i p t i o n i s c h a r a c t e r i s t i c . o f a l l t h e t e r m i t e s w h i c h h a v e d i e d i n t h e 4 t h , 5 t h and 6 t h s e t s .  Observations  made on 4.1.1937.  Table 1# - Measurements i n M i l l i m e t r e s of A l a t e s of Sootermopsis a n g u s t i c o i l i s (Hagen) Maximum Minimum Average Length from t i p to t i p Length of forewings Width of forewings Head l e n g t h Head width Pronotum l e n g t h Pronotum width  29.00 24.50 6.33 2.64 2.64 1.28 2.31  § - Taken from Sumner ('33).  23.50 19.50 5^25 2.20 2.25 1.03 1.94  26.72 22.17 5.81 2.40 2.49 1.17 2.18  X 17.9 4.6 2*12 2.18 0.94 1.90  X  X  2.08 2.42 1.02 1.94  2 o 29 2.48 1.03 2.13  Specimens numbered X are from B r i t i s h Columbia.  Table 2 - Measurements i n M i l l i m e t r e s of^ A l a t e s o f Zootermopsis n e v a d e n s i s (Hagen). L e n g t h f r o m t i p t o t i p21 .0 17 »0 Length of f o r e wings 4 .5 Width of f o r e wings 2 .60 Head l e n g t h 2 . 33 Head W i d t h 0 .94 Pronotum l e n g t h 1 .86 Pronotum w i d t h  22.3 18.0 4.0 2.37 2.22 0.88 1.39  19. 8 16. 0 4. 0 1. 95 2. 04 0. 77 1. 79  22. 5 18. 2 4. 0 2. 16 21 1. 04 1. 86  21. 0 17. 5 4. 2 1. 95 17 0. 94 1. 82  21. 5 18. 0 4. 4 2. 10 2. 27 0. 90 2. 01  21 .0 17 .2 4 .0 2 .08 2 ft £w2 0 .96 1 .93  Table 2 - Continued: L e n g t h f r o m t i p t o t i p20 .7 Length of f o r e wings 17 .0 Width o f f o r e wings 4 .4 Head l e n g t h 2 .04 Head W i d t h 2 .15 0 .97 Pronotum l e n g t h Pronotum w i d t h 1 .87  19.7 24. 0 - 20. 7 16,0 1 9 , 0 16. 8 3.9 4. 8 4. 0 1.93 2. 13 2. 06 1.95 2, 30 2. 18 0,93 1. 09 0, 90 00 1.72 2. 1. 78  21. 7 17. 7 4. 3 2, 00 2. 07 0. 85 1. 82  20. 0 16. 0 4. 0 2. 02 2. 12 0. 82 1. 84  21 ,5 17 .0 4 .0 2 .00 2 .08 1 Qoa 1 .87  Table 2 - c o n c l u d e d : L e n g t h f r o m t i p t o t i p22 .3 Length of f o r e wings 17 .6 Width of f o r e wings 4 .3 Head l e n g t h 2 . 23 2 .09 Head w i d t h Pronotum l e n g t h 1 .02 Pronotum w i d t h 1 *97  22. 3 22.0 17,5 17. 8 4. 0 4.2 2.17 2. 10 2*24 2. 14 1.05 1. 07 1,91 1. 83  21. 3 17. 0 4, 3 16 2 *> 24 1. 02 2 e 00  2 1 . 5x 17. 5 4. 3 1. 90 2. 02 0. 95 1. 85  19 .7 16 .2 4 .0 1, 85 1. 95 0. 91 1. 68  3» 0 18. 4 4. 5 2. 18 2. 37 0. 99 2 » 04 2  A l l s p e c i e s w i t h t h e e x c e p t i o n o f t h a t l a b e l l e d x were t a k e n a t Loughbor© I n l e t , B. C. X was t a k e n a t V a n c o u v e r , B. C.  T a b l e 3- M e a s u r e m e n t s i n M i l l i m e t e r s o f B r a c h y p t e r o u s R e p r o d u c t i v e s of Zooterrnopsis a n g u s t i c o i l i s (Hagen). Length of f o r e wings: Length of h i n d wings: Head l e n g t h : Head w i d t h : Pronotum l e n g t h : Pronotum w i d t h : Length o f body:  T a b l e 4.  3 .0 2. 58 2 .4 2. 27 3 .45 3. 3 2 .84 27 77 1 * 2 _L 1. 1 2 .75 2. 17 14 .5 14. 0  2 .32 1 .78 3 .1 2 .65 1 .25 2 .4 14 .5  Measurements i n M i l l i m e t e r s o f A p t e r o u s R e p r o d u c t i v e s of Zooterrnopsis a n g u s t i c o i l i s (Hagen).  Head l e n g t h Head w i d t h Pronotum l e n g t h Pronotum w i d t h T a b l e 4.  2 .55 3 p 2 .9 1 .82 2 .8 3 .25 2 .35 2 .9 1•1 1 .4 2 .63 2 ,1 13 .0 15 .5  3.14 2.84 1,42 2.48  3.10 '2.58 1.12 2.38  3.3.6 2,88 1.35 2.54  3.38 2.98 -1.26 2.52  3.34 2.80 1.25 2.52  3.52 3.02 1.28 2.52  3.40 2.92 1.28 2.56  3.60 2.82 1,28 2.60  2.82 2.54 1.00 2.18  Continued:  Head l e n g t h Head w i d t h Pronotum l e n g t h Pronotum w i d t h  3.28 3,12 1.25 2.58  3.62 2.82 1.20 2.62  3.60 3.02 1.28 2.86  3.22 2.75 1.15 2.32  Table 5.  Measurements i n M i l l e m e t e r s , and i n d i c e s , of S o l d i e r s of Zooterrnopsis a n g u s t i c o i l i s . (Hagen).  Specimen No. Head l e n g t h - ., Head width * Minimum head width Distance between rims of the antennal foveolae Labrum width Gular l e n g t h ' Maximum gular breadth Minimum g u l a r breadth Pronotum l e n g t h Pronotum width. Mandible l e n g t h Head width-mandible index Head index Head c o n t r a c t i o n index Head-pronotum index Head-mandible index Gular c o n t r a c t i o n index  4.28 4.05 3.18 >.I6 ;.,£o 2.56 0.88 2.94 1.20 0.72 1.52 2.60 2.72 0.67 0.94 0.79 0.64 0.65 0.60  4.78 4.02 3.24  3.90 3,88 3.12  4.00 3.86 3.02  3.68 3.55 2.94  3.76 3.58 2.90  4.14 4.12 3.52  2,62 0.91 3.22 1,34 0.85 1.50 3.18 3.30 0.82 0.84 0,81 0.79 0.69 0.63  2.50 0.85 2.42 1.18 0.80 1.55 3.00 3,38 0.88 0.99 0.81 0,77 0.87 0.68 X  2.68 0.90 2.75 1.20 0.82 1.40 2* 2 3,30 0,86 0.96 0.78 0.70 0.83 0.68 X  2«5 <» 0.94 2.90 1.10 0.70 1.40 2.82 3.14 0.88 0.97 0.83 , 0,80 0.85 0.64 X  2.52 0.94 2,98 1.10 0.68 1,35 2.86 3.22 0.90 0,95 0.81 0.80 0.85 0.62 x  2.85 0.96 3.12 1,46 0,68 1.60 3.12 3.54 0.86 1.00 0.85 0.76 0.85 0.47 X  4 .18 3 .68 3.10  4.10 3.54 3,00  1001 3.06 3.10 2.60  A 1002 4.50 3.45 3,00  B 1002 4.47 3.48 2,. 86  A 1003 4.62 3.57 2.98  2 0 .98 2 .93 1 .24 0 .70 1 .50 3 .00 3 .00 0 .81 0 .88 0 .84 0 .81 0 .72 0 .57  2.30 0,88 2.74 1.08 0.60 1.40 2.68 3.08 0.87 0.86 0.85 0.76 0.75 0,55  3.08 0.92 3.48 1.16 0.62 1U25 * 52 3.60 1.04 0.77 0.87 0,73 0.80 0.54 X  2.80 0.92 3.66 1.03 0.55 1.30 2 ,78 3.68 1.06 0.78 0.82 0.80 0.82 0.53 X  3.03 0.92 3.68 1.10 0.60 1.40 2.77 3,65 1.02 0.77 0.84 0.77 0.79 0.55 X  Table 5. - Continued: Specimen Ho. Head l e n g t h Head width Minimum head width Distance between rims of the antennal foveolae Labrum width Gular l e n g t h Maximum g u l a r . b r e a d t h Minimum gular breadth Pronotum l e n g t h Pronotum -width Mandible l e n g t h Head width-mandible index Head index . • Head c o n t r a c t i o n index •Head-pronotum index Head-mandible index Gular c o n t r a c t i o n index  4.48 3.78 2.90 2.72 0.88 2.88 1.00 0.68 1.56 2.90 3.10 0.82 0.84 0.77 0.78 0.69 0.68  (  2.44 0,98 2,28 1.00 0.76 1,22 2.52 2 » *3 6 0.83 1.01 0.84 0.81 0.84 0.76 X  Table"5. - C o n t i n u e d : Specimen Ho. Head l e n g t h Head w i d t h ' Minimum'head'width D i s t a n c e between rims o f the a n t e n n a l foveolae Labrum w i d t h Gular Length Maximum g u l a r b r e a d t h Minimum g u l a r b r e a d t h pronotum l e n g t h Pronotum w i d t h Mandible l e n g t h Head width-mandit> 1 e i n d e x Head i n d e x Head c o n t r a c t i o n i n d e x Head-pronotum i n d e x . Head-mandible i n d e x Gular c o n t r a c t i o n i n d e x ;  B 1003 4.73 3.82 3.28 3.18 0.98 3.26 1.14 0.78 1.42 3.00 3.18 0.83 0.81 0.86 0.78 0.67 0.68  1004 4.27 3.95 3.05  1005 3.96 3.13 2.56  1006 4.48 3.47 2.98  1007 5.38 4.32 3.56  A 1008 4.18 3.72 3.18  B 1008 4.07 3.56 3.00  3.23 0.80 3.00 1.18 0.78  2.60 0.52 2.68 1.00 0.54 1.20 2.42 2.85 0.91 0.79 0,82 0.77 0.72 0.54  2.87 0.80 3.30 1.12 0.60 1.26 2.91 3.50 1.01 0.77 0.86 0.84 0.78 0,54  3.40 0.68 4.02 1.22 0.70 1.66 3,46 3,65 0.84 0.80 0.83 0.80 0.68 0.57  3.24 0.90 2.98 1*30 0 .82 1.24 2.80 3.30 0.89 0.89 0.85 0*75 0.79 0.63  3.03 0.82 3.00 1.16 0.68 1.25 2.73 3.18 0.89 0.87 0.84 0.77 0.78 0.58  4.,73 4,,10 3,.10  4v 50 3. 83 3. 10  3..29  o, 32  0 .77 3 .50 1 .16 0 .68 1 .57 3 .09 3 .74 0 .91 0 .87 0 .76 0 .75 0 .79 0 .59  1. 05 3. 07 1. 18 Q. 58 1. 40 2 «66 3. 14 0. 82 0.,85 0..81 0,,70 0,,70 0,,49  3.24 0.82 0.92 0.77 0.76 0.66  X  T a b l e 5* - C o n c l u d e d : Specimen Ho * Head L e n g t h Head w i d t h Minimum h e a d w i d t h D i s t a n c e between rims o f the a n t e n n a l foveolae Labrum w i d t h Gular l e n g t h Maximum g u l a r b r e a d t h Minimum g u l a r b r e a d t h Pronotum l e n g t h Pronotum w i d t h Mandible l e n g t h Head w i d t h - m a n d i b l e i n d e x Head i n d e x Head c o n t r a c t i o n i n d e x Head-pronotum i n d e x Head-mandible i n d e x Gular C o n t r a c t i o n index  1009 4,,44 4,.08 3-,»2 2  1010 4..32 3,.75 3 ,48  3 ,37 1 .00 3 .15 1 .30 0 .72 1 .47 2 .76 3 .40 0 .83 0 ,92 0 .79 0 .68 0 .77 0 .55  3 .38 0 .73 3 .42 1 .36. 0 .90 3 .30 0 .88 0 .87 0 ,93 0 .76 0 .66  X  4,.03 3..50 2,,56  y 4;.28 3. 35 2. 42  2 .70 0 .70 3 .10 1 .08 0 .48 1 .53 2 .48 3 .19 0 .91 0 .87 0 .73 0 .71 0 .79 0 .44  2. 73 0. 81 3, 28 1* 25 0. 73 1. 38 2. 70 3. 26 0. 97 0, 78 0. 72 0..81 0..76 0..58  S p e c i m e n s x a n d y a r e Z. n e v a d e n s i s California respectively.  t a k e n i n Or eg'  PLATE I.  1 and 2. D o r s a l  a n d v e n t r a l v i e w s of t h e head o f an a l a t e  of Z o o t e r i r i o n s i s a n g u s t i c o i l i s . x 1 8. 3 a n d 4 . Same v i e w s o f a l a t e R e t i c u l i t e r m e s  hesperus. x 7 o .  5 and 6. Same v i e w s , o f n e w l y h a t c h e d nymph of Z o o t e r r n o p s i s a n g u s t i c o i l i s . x90.  PLATL I  PLATS I I . 1.  S i d e viev; of s o l d i e r  of Z. a n g u s t i c o i l i s . x 1 1 . 3  2.  S i d e viev; o f s o l d i e r o f J3L. i i g ^ p a i l l i s . . x 2 2 .  3.  Labruin o f s o l d i e r  o f Z. n e v a d e n s i s ,  showing  tormae.  x37.  4. Labrum o f a l a t e  o f Z. a n g u s t i c o i l i s , v e n t r a l  aspect,  s h o w i n g e p i p h a r y n x and t o r m a e . x 3 7 . 5. Labrum o f a l a t e  of R. h e s p e r u s . x 3 7 .  6.  Labrum o f s o l d i e r  o f Z. a n g u s t i c o i l i s . x 3 7 -  7-  Labrum o f s o l d i e r  o f R. h e s p e r u s . x 8 8 .  8.  Lateral  a s p e c t o f h y p o p h a r y n x o f s o l d i e r o f Z,  angusticoilis. 9. D o r s a l a s p e c t of sume.  P L A T E II  PLATE I I I . 1. D o r s a l a s p e c t of m a n d i b l e s 2. V e n t r a l a s p e c t o f m a n d i b l e s 3.  D o r s a l a s p e c t of mandibles  of a l a t e  of Z.  of a l a t e  angusticoliis.  o f R.  hesperus.  x'76.  of s o l d i e r  of Z.  angusticoliis.  4. D o r s a l a s p e c t o f m a n d i b l e s  of s o l d i e r  o f R.  hesperus.  5. D o r s a l a s p e c t o f m a n d i b l e s  of s o l d i e r  of Z.  nevadensis.  x  37.  X18. 6. D o r s a l a s p e c t o f m a n d i b l e s Z.  angusticoliis.  x183.  of f i r s t  i n s t a r nymphs o f  P L A T E 111  I  0  PLATS Phylogenetic  IV.  t r e e of I s o p t e r a . A f t e r H a r e  '37-  PLATE. W  PLATE V.  1. V e n t r a l  a s p e c t of r i g h t m a x i l l a o f an a l a t e o f Z,  angusticoliis.  z.37•  2.  D o r s a l a s p e c t o f same. x 3 7 .  3.  D o r s a l a s p e c t o f l e f t m a x i l l a • • o f a n a l a t e o f R. hesoerus. x88.  4. V e n t r a l  a s p e c t o f same. x 8 8 .  5. D o r s a l a s p e c t o f m a x i l l a o f a s o l d i e r o f Z. a n g u s t i c o l i i s . x24. 6.  Ventral  a s p e c t o f m a x i l l a o f a s o l d i e r of R. hes e r a s .  x37»  PLATE VI.  1. L a b i u m o f a s o l d i e r  o f Z. a n g u s t i c o i l i s ,  ventral  X18. 2. Same o f R. h e s p e r u s . x 4 6 . 3. T e n t o r i u m o f an a l a t e  o f R. h e s p e r u s . x 4 6 .  4. Same o f Z. a n g u s t i c o i l i s . x 1 8 . 5. T e n t o r i u m o f a s o l d i e r  o f Z. a n g u s t i c o i l i s  6. Same o f R. h e s p e r u s . x 4 6 .  • X18.  aspect.  P L A T L VI  PLATS V I I . 1 . T e n t o r i u m of 'brachypterous r e p r o d u c t i v e  o f Z.  a n g u s t i c o l i i s . x18. 2. T e n t o r i u m o f f i r s t  i n s t a r nymph o f Z. a n g u s t i c o l i i s . x 1 0 9 .  3. T e n t o r i u m o f s e c o n d i n s t a r 4. T e n t o r i u m o f t h i r d i n s t a r  nymph. nymph.  P L A T E V\l  PLATE  VIII.  L a t e r a l a s p e c t o f c e r v i c a l r e g i o n and p r o t h o r a x o f a n a l a t e o f Z. a n g u s t i c o i l i s . x 2 0 . V e n t r a l a s p e c t o f same. x 2 0 . A diagram of  showing the r e l a t i v e  s i z e s of the p r o n o t a  t h e c a s t e s o f Z. a n g u s t i c o i l i s . x 3 7 .  • 1.Brachypterous s o l d i e r . 2. Typical soldier. 3 . Apterous r e p r o d u c t i v e . 4. A l a t e . 5. B r a c h y p t e r o u s r e p r o d u c t i v e . The p r o n o t a o f a s o l d i e r  and an a l a t e of Z. n e v a d e n s i s .  PLATE 'IX 1. P r o n o t a of Z.  angusticoliis.  1. L o n g b r a c h y p t e r o u s 2. S h o r t » 3. A l a t e . 2. P r o n o t a of R.  hesperus.  1. A l a t e o f Z. 2. A l a t e . 3. S o l d i e r . 3- L e f t s i d e  Same w i t h  re-productive. ' I"  x88.  angusticoliis.  of t h e m e s o t h o r a x o f an a l a t e of  angusticollis, 4.  x37.  the wing  Z.  s h o w i n g a x i l l a r y s c l e r i t e s . x37«  expanded.  x37.  P L A T E IX  PLATE X. 1. K e t a t h o r a x o f an a l a t e  o f Z. a n g u s t i c o i l i s . x 3 7 .  2. N o t a o f a n a l a t e o f R. h e s p e r u s . x 4 0 . 3. M e s o t h o r a x  o f an a l a t e  o f R. h & s p e r u s . x l 8 5 .  4. M e t a t h o r a x o f a n a l a t e o f R. h e s p e r u s . x 1 8 5 . 5. P l e u r a o f m e s o t h o r a x 6. Same o f R. h e s p e r u s .  o f a n a l a t e o f Z. a n g u s t i c o i l i s . .  PLATE X  PLATE XI. 1. S t e r n a oi' an  a l a t e of Z.  2.  Same of R.  hesperus.  3.  L a t e r a l a s p e c t of t h e a l a t e of > plates  4.  Z.  angusticoliis, dorsal  x46. v e n t r a l surface  a n g u s t i c o l i i s , showing  I n the m e s o t h o r a x and  C a u d a l v i e w of t h e  of t h e  n o t a of  strengthening  metathorax.  anterior portion  showing e n d o s k e l e t a l  structures  of  prothorax,  of an  alate  Z. a n g u s t i c o l i i s . 5.  M e s o t h o r a c i c p l e u r a l apodeme of an angusticoliis.  aspect.  alate  of  Z.  of  an  P L A T E XI  o  PLATE X I I . 1 . Fragmata  o f meso and m e t a t h o r a x  and f i r s t  abdominal  segment o f a n a l a t e o f Z. a n g u s t i c o i l i s • x 2 5 . 2. Same o f R. h e s p e r u s . x 4 0 . 3.  Sternal endoskeletal reproductive  4. Same of ,a s o l d i e r  structures  o f Z.  of a brachypterous  angusticoilis.  o f R. hesr>erus. X 1 3 0 .  5. The l e g s  o f a n a l a t e o f Z. a n g u s t i c o i l i s . x 1 4 .  6.  o f an a l a t e o f R. h e s p e r u s . x 3 7 .  The l e g s  P L A T E XII  PLATE X I I I .  1. M e t a t h o r a c i c  l e g o f a s o l d i e r o f Z. a n g u s t i c o l i i s . x 1 3 -  2.  Same o f a f i r s t  3.  L e g s o f a. s o l d . i e r o f R. h e s p e r u s . x 3 7 .  4.  Tibial  spurs  i n s t a r nymph o f Z. a n g u s t i c o l i i s . 4 3 .  o f an a l a t e  5. P r e t a r s u s o f an a l a t e  o f Z. a n g u s t i c o l i i s .  o f Z.  angusticoliis.  6. Same o f a s o l d i e r . 7.  Wings o f Z. a n g u s t i c o l i i s . x 2 . 5 .  S. \Ting o f R . h e s p e r u s . x 8 .  P L A T E X\U  PLATS XIV. 1. B a s a l p o r t i o n o f w i n g o f an a l a t e o f Z. 2.  angusticollis  Same o f I . ' e t a t h o r a c i c w i n g .  3. and4. YJings o f s e v e n t h i n s t a r nymph, Z,  angusticoliis  5. Same o f R. h e s p e r u s • 6. Vfing p a d o f an e i g h t h i n s t a r " b r a c h y p t e r o u s r e p r o ductive  o f Z. a n g u s t i c o l i i s .  7.Same o f s e v e n t h i n s t a r nymph.' 8. Same o f l o n g b r a c h y p t e r o u s 9. Same o f a s h o r t  "  reproductive. "  P L A T E X\V  FLAffiS  1.and  XV.  2. Wing d e v e l o p m e n t o f a p t e r o u s r e p r o d u c t i v e s Z. c  angusticoilis.  3 and 4. Same o f ' . s o l d i e r s o f Z,  angusticoilis.  of  P L A T E . XV  2  PLATE XVI. 1. S t e r n i t e s  o f f e m a l e abdomen o f an a l a t e o f Z . a n g u s t i c o l i i s  2. Same o f a m a l e . 3,4 and 5- D e t a i l s  of female g e n i t a l i a .  P L A T E XVI  PLATE XVII.  A l a t e Zooterrnopsis a n g u s t i c o i l i s . X7.2. 1. D i g e s t i v e  tract.  2. Female r e p r o d u c t i v e  system.  3. M a l e r e p r o d u c t i v e s y s t e m . 4. Hervous 5.  Heart.  system.  P L A T E XVII  V  4  5  PLATE X V I I I .  1. B r a i n o f a l a t e  Zootermopsis a n g u s t i c o l i i s  s t o m o d a e a l n e r v o u s system,, x45. 2.  S e c t i o n of a l a t e  brain  Zootermopsis  Semidia.gr ammat i c . angusticoliis  through suboesophageal g a n g l i o n . 3. L a t e r a l  a s p e c t of b r a i n  Diagrammatic.  showing  a t the median  x45. line.  P L A T E XVIll  PLATE X I X . 1.  Streblomaatix  strix.  x15C0.  2. Dlnenyrrroha f i m b r i a t a . 3. Pryaonympha  x700.  m a j o r , x55D.  4. H e x a m a s t i x t e r m o p s i d i s .  x2500.  5.  Tricercomitus  termopsidis.  6.  Trichomonas t e r m o p s i d i s . x  x200o.  15°0«  7. Holomaa t i g o t e s e long, a turn. x5°0. S • Sp i r o t r i chonympha f l a g e l l a t a . 9• S p i r o n y m p h a p o r t e r i .  x  10 . Torquenyiapha o c t o p l u s . x 1 8 0 0 . 11.  Trichonympha c o l l a r i s .  12.  T.  campanula. X 1 7 0 .  13.  T.  sphaerlca.x 1 7 0 .  14. T. a p ; i l l s .  x660.  x 170.  x 7 0 0.  P L A T E XIX  PLATE  Associations  of H i r m o c y s t i s  XX.  termitis.  x109.  PLATE. XX  

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