Proceedings, 1936 25

A PRELIMINARY LIST OF IN BRITISH COLUMBIA

13} J. K. J acoh \ Ta ncouvel'. B.C.

The presence of protozoa in t he 10\\'er term ites has long been kno \\'n and ,,'as fir st recorded by L espes in 1856 , Sin ce then. m any genera and species haH been cli scO\'erecl, The opinion formerly held by som e en­ tom ologists was that there \\'as som e correlation exi:; ting bet\\'een the presence o f protozoa and the condition of the sexua l organs. ]~ece nt researches w ould indicate t hat the majority of protozoa are sy mhiotic and that their presence ha:; nothing to do with the sexua l organs, To get a clear understanding of the relati onships of protozoa and termites. it is necessary to present a brief description of t he dige:; ti,'e tract of those which contain these symbionts. E mbryonically, the gut is divisible into three \\'ell-defined regions ; the fo re-intestine or stomodeum, the mid-intestine, and the hind-intes­ tine or proctodeum. The stom odeum a nd the proctodeum are f.o rmed embryonically as in vaginations of the anterior and posterior body wall respectiyely. The lining. therefore. o f the fore- and hind-intestine is of t he sam e material as that ·o f the exoskeleton, that is. chitin. The mid­ inte,; tine ari ses independently of the hody wall and thus lacks this chitinous lining, The thin-walled pharynx m erges into the oesophagus and a large crop. follo wed by the gizzard. P osteri or to the g izzard the foregut protrude:; into the cavity of the mid-intestin e, f.orming a large oeso­ phageal yah'e, At the anterior end of the mid-intestine. \\,hich i:; tubular and of uniform size throug hout, are four short enteri c caeca. At the junction of the mid- and hind-inte:; tines are a number of ;\[alphigian tuhules. Posterior to these tubules. and in the enlarged portion o f the hind-in testine, are present large numbers of protozoa. T he larger m em­ hers of this protozoan population appear to li,'e in a symbioti c relation­ ship with their host, In return fo r food. protection and a :i uitable medium in which to li ,'e. they render the a sen 'ice in the dige:;­ ti on of cellulose which the termite is unable to perform alone, According to Beall's (2) and my own coll ection the foll o ll'ing are the three species of termite:; in British Columbia:- Zootermopsis angusticollis ( Hag en ) . Z. nevadensis (Hagen ) and Reticulitermes hesperus (Banks). In the fi rst two of these specie s the d igesti I'e tract is a s described above. The protozoa occurring in these are as foll ows. those I have found being indicated by an asterisk : Z. angusticollis, the common damp-wood termite. 26 B. C. Entomological Society

* Trichonympha collaris (Kirby), * T. campanula (Kofoid and Swezy), * T. sphaerica (K o foid and Swezy) , * Streblomastix strix (Kofoid and S\\'ezy), * Trichomonas termopsidis (Cle\'eland ) , * Tricercomitus termopsidis (Kirby) , * Hexamastix termopsidis (Kirby), * Hirmocystis termitis (Leidy), Z. nevadensis, the small clamp-wood tennite. is heli e\'ed to haye the sam e spec ies 'of protozoa as Z. angusticollis but as yet I ha \'e not ex­ amined any o f t his species, R. hesperus (Banks) . the western subterranean termite, * Trichonympha agilis (Leid y), Microjoenia ratcliffei (Brown ) , * Torquenympha octoplus (Brown ), * Spironympha porteri (Koid zu1l1 i) . S. ovalis. * Spirotrichonympha ftagellata (G rassi and Foa). * Holomastigotes elongatum (G ra ~s i ), Dinenympha fimbriata (Kirby) Dinenympha sp, * Pyrsonympha major (Powell ), P. minor. P. granulata. * Hirmocystis termitis ( L eid y), P rotozoa are not jJ l'esent in ne\\'ly-hatched termites. T hey are feci for some time on materi a l (stomodeal food) regurg itated by t he parents but after a m oult or t wo. proctodeal food is added and finall y con­ st itutes almost the entire food supply procured from another indi\idua l. As there are no protozoa in the fore-intestine of the parents. fi rst in stal' term ites recei\'e no protozoa in stom odeal food , T here are protozoa in a ll the old er in stars, Nymphs. soldiers and adults conta in t he "am e s pecies of protozoa. b ut they d iff er in total population and possibly relati \'e proportion in t he different forms of termites, :\ dults contain felVer protozoa than cl o ny mphs o f the se\'enth in star. T he total num­ ber of protozoa found in immature termites \'aries \I'ith the time since the last m oult and the nearness to t he next m oult, A m onth before t he m oult. the intestin e appears normal in el'er)' way and possesses its us ual content o f food m aterial and protozoa, F rom three to four \\'eeks before the m oult, the term ite enter s a peri od of self-stan 'ation. so t hat the food content of the in testine disa ppears but the protozoa rema in , H owever, with t heir food su pply cut off, the wood-eating protozoa gradually stan'e and disappear. Of these, the three species of Trichonympha g o fi rst foll owed by Trichomonas, so that Proceedings, 1936 27 all four are completely absent before the moult occurs. O f the other three species of protozoa present. Streblomastix may, or may not remain throug h the peri od of the moult and Tricercomitus a nd Hexamastix always persist in the intestine. A s was said before, the fo re-intestine a nd hind-intestine ,,'a ll s are lined with chitin w hi ch is continuous. throug h the m outh and anu,.;. ,,·ith the exoskeleton o f the in sect; ,,· ith the shedding of the exoskeleton. t hese chitinous linings are also shed. Ha\"ing thus renovated its di ge,; ti\'e tract. the termite must reacq uire a protozoan fauna. The ah(we three sma ll er. non-symbioti c spec ies o f protozoa m ay rem ain in the g ut throug h the m oul t; the larger specie,; must he reacquired from other termites. T he exact m ethod o f refaunation is not definit ely known. That refa unation does not take place by anal contact w ith protozoa or faeces is shown by the careful studies o f A ndrew ( 1). l{cfaunatio n does take place readil y hy feedin g on m ateri a l containing protozoa. T he m ethods by which the protozoa can be normally obtained for this feed­ in g \\"e re not definitel y determined in Andre\\"s experiments. Faeces do not contain protozoa in m otil e condition or in cysts. thoug h there is some indicati on that cystoid form s of Trichomonas termopsidis and Trichonyrn,pha campanu1a exist in the intestine under certain unfa\'orable conditions. Feeding term ites on faeces does not refaunate them. Can­ nibali sm leads to a transfer of protozoa but it is not the onl y agency that achie\" es t hi s result. P rotozoa have been found in som e in stances on the un extruded pe ll et and in the first drop of intestinal fluid . hen ce the transfer of protozoa during proctodeal feeding may be one of the normal m ethods of refaunation. A ndrew states that in mixed groups of faunates and defaunates \\'here canniba li sm \\'as pre\'entcd and no stom odeal fe ed­ in g or dropping o f liquids \\'as observed, proctodea 1 feeding on the part of both faunates a nd defaull ates continued as usua l, but refaunatioll \\"as not accom pli shed. indicating that this m ay not be the m ost effecti\'e m ethod of refaunation. In a few cases. liquid droppings were found to contain li\ing fl agell ates, so that fe eding on the,.;e and on fresh n e,; t structures in \\' hi ch intestinal fluid is used as a cem ent Illay probably account in part for normal refaunatio ns. By usin g recently m oul ted indi\'iduals it \\'a s determined that in the normal colony . refa ull ation has usually taken place by the t hird or fourth cl ay. Careful studies of the number of protozoa in a large nurmal ny1l1ph ha \'e sho \\'n approximately 25.000 Trichonymphs and ahout 500.000 Trichomonas ( 1) . Gregarines are a lso present in Z. angusticollis at least . and probably in R. hesperus. These are purely parasitic and are located in the anterior part of the mid-intestine. Their population vari es in a sing le indi\'idua l from one or 1\\'0 to ,;e \'eral hundred (8) . In the clevelo pment, the im­ mature g regarines are fir st attached to the intestinal wall in th e crc\'ices 28 B. C. Entomological Society between tvvo g roups of cell s. Later the epimerite of the young gregarine is l o~t and then the protozoa becomes free. These parasites have' no apparent effect u pon the ceIL:; of the in testine, nor is t he termite y isibly affected by the pre:;ence of large numbers of gregarine ,.; . Hirmocystis termitis i:; the C0111 111 0n species I hal'e fo und in Z. angusticollis. Kofo­ idina ovata (Benry). a second g regarine. is recorded in Z. angusticollis fr0 111 t he S tate of vVashingt on. ,\s yet. th is species has not been fo und in a ny of the B ritish Columbia specimen s. Spirochaetes and bacteri a are fairly common in the inte:; tine. Some :;pirochaete,.; a nd a re normally fo und adherent to the surface of the flagel lates. Some of the Aagellates hale in terna l para,;ites of the ir O\l"n , generally bacteria. F rom this work, it can be seen that the protozoa in British Columbia t ermites a re silllila r to those exi,; ting in s imilar termite,; of Califo rnia.

Bibliography

1. ,\ ndrcII' . B . .1 .- 1930. l\Iethod and rate of protozoa refaunation in the termite. Termopsis angusticollis. L' nil·. Cali f. P u bl. Zool. . 33: 449-470. 2" Beall . C.-193 1. Notes on the terl11 ites of I3ritish Cul umb ia, Proc. E nt. Soc. 13r. Col. . 28 : 33-35. 3. B rown. V . E.- 1930. Hyperlll astigote Aagell a t es from the t ermite R ecticulitermes. Torquenympha octoplus gen. n OI', s p. no\'. a nd hl"O new specie,; of Microjoenia. U nil", Cali f. P ubl. Zoo1. 36 :67-80. 4a. Cleveland, L. R.-1923. Correlation between the food and mor­ phology of termites and the pre,;e nce of intestinal protozoa. Amer. J ourn. Hyg. 3 : 444-46 1. h. 1925. The effects of oxygenation and stan"ation o n the bctl\"ee n the termi te Termopsis and its intestinal Aagella tes. B io I. Bull. 4R: 309-326· c. 1925. The feeding habit of t ermi te castes a nd its relatio n t o their intestinal flagellates. BioI. Bull. 48: 295-308. Sa. Heath. H.- 1903. The habit,; of California terl11ites. B ioI. Bull. 4: 47-63. b. 1927. Caste formation in the termite genus Termopsis. Journ. Morph. Physiol 43: 387-424. 6. Heath , H. and W ilbur, B. C.-1927. The development of the soldier caste in the termite genus Termopsis. BioI. B ull. 53: 145-150. 7. ] 1111ll S, , \ . D.-I <)25 . 1\ General Textbook of Entomc logy. ( L o n­ do n. Methuen). 8 Henry. Dora. P.- 1933. Hirmocystis termltls (Leidy ) and Kofoid­ ina ovata gen. 1101", sp. n OI·. from termites. Arch . . 80: 101- 11 5. Proceedings, 1936 29

9. K icki er. C. \\'.-1929. Streblomastix strix, morphology and Illito,; is. C ni \'. Calif. Pub!. Zoo!. 3.3: 109- 122. l Oa. I(irby . H.-1923. -;\I orphology and lllito,; is of Dinenympha fim­ briata ,.; p. no \·. C ni \'. Cali f. P ub!. Zool. 26 : 199-220. h. 1930. T ri chomonad Aa gell ates fr om termites IT Tricercomitus gen. no\·. and Hexamastix ,-\ Iexeieff. L'n i\'. Calif. P ubl. Zou!. 33: 39.3-4-1-4 . c. 193 1· Trichom onad Aagell ates from termites IT Eutrichomastix and the ,.; ubi