ANNALES DE LIMNOLOGIE, t. 3, fasc. 2, 1967 : p. 231-247

D E S C R I P T I O N S O F T H E L A R V A E

O F ATURUS SGABER KRAMER,

PROTZIA EXIMIA PROTZ

A N D PIONA UNCATA K O E N I K E

W I T H N O T E S O N T H E L I F E - H I S T O R I E S

O F T H E L A T T E R T W O .

[Hydracar ina]

by R . K . H . JONES.

Al though m o r e t h a n 3 000 species of wa t e r -mi t e s have been descr ibed, the l ife-histories a re k n o w n for less t h a n 5 % of t h a t n u m b e r , and compara t ive ly few juvén i le s tages have been descr ibed. Th i s p a p e r descr ibes t h e l a rvae of th ree c o m m o n mi tes , one of w h i c h w a s prev ious ly widely believed no t to p rodnce larvae. Many mi tes rwith su r f ace - runn ing la rvae have been observed bu t the préc ise me thod by w h i c h thèse la rvae a t t a in a pos i t ion on the i r hos t s h a s been a m a t t e r of con jec ture . T h e me thod is described for Protzia eximia PROTZ wh ich also provides the first publ i shed example of a mi te whose la rvae pa ras i t i se adul t Caddis Fl ies .

As l i fe-history records a re d ispersed t h r o u g h o u t a cons idérable and diffuse l i t e ra tu re , it is t h o u g h t dés i rab le to b r ing toge ther in one table p a r t i c u l a r s of the 153 mi tes w h o s e l ife-histories have been publ ished.

1. — GENERAL N O T E S .

1.1. — A tu rus seaber KRAMER.

Thi s mi te lives in fast-flowing w a t e r and t h e females lay one or two eggs only a t one t ime . T h e eggs develop slowly wi th in the females and it h a s the re fore been a s sumed by m a n y a u t h o r s t h a t t hey would p robab ly h a t c h d i rec t ly to n y m p h s . In o rde r to tes t th i s idea I careful ly isolated eleven ovigerous females un t i l t hey p roduced , in ail, 12 eggs, f rom w h i c h 12 la rvae eventua l ly ha t ched . T h e l a rvae lived for abou t t h r ee weeks in the l abora to ry bu t n o n e formed n y m p h o c h r y s a l i d s , suggest ing t h a t a pa ras i t i c

Article available at http://www.limnology-journal.org or http://dx.doi.org/10.1051/limn/1967011

2 3 2 R. K. H . JONES (2) stage m a y be needed. Larvae were subsequen t ly found amongs t mosses in r ivers du r ing the m o n t h s of J u n e and Ju ly , bu t none h a s so far been found a t t ached to a hos t .

1 . 2 . — Piona uncata KOENIKE.

Adul t females of th is species a re found in p l a n k t o n of ponds and di tches as ear ly as March 1 st and are usua l ly c o m m o n by ear ly Apri l , r e m a i n i n g so unt i l J u n e . In the l abora to ry they ca tch and feed on Cladocera and th i s seems to be the i r m a i n diet . Eggs a re laid in the l abora to ry a lmos t a lways w i t h i n a week of the females ' c ap tu re , bu t wild females a re a lways full of eggs up to the first week in May, so tha t it is probable t h a t the n o r m a l egg laying per iod outs ide is in May and tha t it is the rise in t e m p é r a t u r e in the l abora to ry t h a t p romûtes t he ear l ier egg-laying. The eggs a re laid in long s t r ings con ta in ing 6 - 4 0 eggs. The s t r ings a re usua l ly a t t ached to algal t h r e a d s or to pond weeds . In the labora to ry the larvae h a t c h in two to th ree weeks and will live for 4 - 6 weeks . STOUT [ 1 9 5 3 b] found t h a t larvae of the subspecies P. uncata exigia V I E T S developed to n y m p h s in 2 weeks w i t h o u t pa ra s i t i sm, bu t m y spéc imens showed no signs of m e t a m o r p h o s i s . In the field the la rvae w e r e in fact found to be paras i t i c on the midge Chironomus plumosus L. and o the r Chi ronomids . T h e i r m o u t h p a r t s w e r e inser ted in to the a r th rod ia l m e m b r a n e s of the hos t ' s abdomen and as m a n y as 2 0 mi te la rvae were found on a single host . T h e hos t midges were kep t in glass tubes con ta in ing a pièce of ru sh s t and ing in a few c m 3 of wate r . T h e mi tes , on leaving the i r hos t s , m a d e the i r way down into t he wa te r and formed the i r n y m p h o c h r y s a l i d s e i ther on the bo t tom of the tube or in the p i th of t he ru sh , often c rawl ing 1 cm into the p i th before set t l ing down. T h e n y m p h s ha tched in th ree w e e k s and formed the i r te le iochry-sal ids abou t th ree m o n t h s la ter .

It h a s been repor ted by CROWELL [ 1 9 6 0 ] for Thyas stolli KŒNIKE, BÔTTGER [ 1 9 6 2 ] for Piona nodata MULLER and PIERSIG [ 1 9 0 0 ]

and CROWELL [ 1 9 6 0 ] for Limnesia undulata MULLER t h a t thèse

mites have a l t e rna t ive m e t h o d s of development , large eggs giving rise d i rec t ly to n y m p h s whi le smal le r eggs p roduce la rvae w h i c h m u s t unde rgo a pa ras i t i c p h a s e before reach ing the n y m p h a l s ta te , and the s i tua t ion in P. uncata KOENIKE m a y well be s imi lar .

1 . 3 . — Protzia eximia PROTZ.

T h e eggs of th i s species w e r e found t h r o u g h o u t t he w i n t e r in crevices in boulders in fast flowing r ivers . Egg ba tches w e r e collected and divided in to th ree sepa ra te tubes , one of w h i c h w a s kept in the d a s h at room t e m p é r a t u r e whi le the two o thers were placed in a ref r igera tor at 5 , 5 ° C. The eggs of the first ba tch ha tched

(3 ) LARVAE OF HYDRACARINA 233

after 18 - 23 days and the la rvae lived for abou t th ree w e e k s . T h e second ba tch w e r e raised to room t e m p é r a t u r e on the 23 rd day and ha t ched in a fu r the r 18 days , at w h i c h t ime the th i rd ba t ch w a s removed from the ref r igera tor and also ha tched af ter a fu r the r 18 days . Th i s seems to offer conclusive évidence t h a t t e m p é r a t u r e is a cr i t ical factor in the h a t c h i n g of thèse eggs.

T h e la rvae pass t h r o u g h the w a t e r film and r u n act ively on the sur face . About two days after h a t c h i n g they are able to j u m p a d i s t ance of nea r ly 2 cm from the w a t e r surface . T h e y are found in the s u m m e r m o n t h s in large n u m b e r s on the moss of bou lde r s wh ich p r o t r u d e from the wa te r . It h a s been a s sumed by m a n y a u t h o r s t h a t j u m p i n g hab i t observed in some H y d r a c a r i n e larvae was associated w i t h the a t t a i n m e n t of a posi t ion on the hos t and observat ion of th i s species proved th is to be so. It w a s found t h a t w h e n the res t ing la rvae a re wa lked over by the fly Weidemannia bistigmata CURTIS or by a n unidentif ied female Caddis fly, t he larvae j u m p e d u p w a r d s , l and ing on the insect , and rap id ly adop t ing a pa ras i t i c a t t a c h m e n t to it. Pa ras i t i sed hos t s were found in consi­dérable n u m b e r s from ear ly May un t i l September . Th i s can be t aken as évidence of a second egg-laying per iod, as it is un l ike ly , in view of the l abora to ry h a t c h i n g tes ts , t h a t larvae of the w i n t e r eggs wou ld h a t c h so late in s u m m e r .

Of the hos t insects c augh t and examined , in May 45 ou t of 90 were paras i t i sed by a tota l of 103 mi te larvae , whi le in Sep tember of 24 hos t s examined 15 w e r e paras i t i sed by 23 mi te larvae .

2. — LARVAE.

T h e n o m e n c l a t u r e used for the br is t les is t h a t proposed by SPARING [1959] p . 13, " S c h é m a der K o r p e r b e b o r s t u n g ".

2 .1 . — Aturus scaber KRAMER.

The larva is oval in out l ine wi th a d is t inc t and deep invagina t ion in the pos te r io r end, r each ing fo rwards fu r the r t h a n the r e a r edges of the dorsa l and ven t ra l p la tes . T h e excre tory fleld is w i t h o u t a p la te but two pa i r s of long spines ( represent ing p robab ly Exp . 1 and Exp . 2) lie close to the excre tory pore . Ex . 2 is inser ted on the edge of the th i rd ep imeron . Ex. 4 is very long and inser ted on a p r o t u b é r a n c e s imi lar to t h a t in Hygrobates spp. P . 3 bea r s a powerfu l curved spine. P . 4 ends in a s t rong a lmos t s t r a igh t c law and bea r s two br is t les . P . 5 is very smal l and bea r s a shor t curved spine .

T h e dorsal br is t les a re very long, P d b . 1 reach ing 55 y. and Db. 1 61 /x. T h e s u t u r e be tween E. II and E. III is p la in ly visible

234 R. K. H. JONES ( 4 )

F I G . 1 : Aturus scaber K R A M E R , larva , dorsa l surface . — F i e 2 : ld° , ven tra l surface .

as far as t h e r e a r m o s t po in t of the E. I and is a lmos t para l le l to the body axis . T h e br i s t le E p . 4 is inser ted some d i s t ance f rom the s u t u r e .

M e a s u r e m e n t s (ju) : 1 2 3

Lengh t of body 230 225 225 Bread th 140 140 140 Cap i tu lum 55 55 55 Pos te r io r br is t les 115 110 110 1 st leg 140 135 140 2 n d leg 140 135 145 3 rd leg 160 140 160

2.2. — Piona uncata KOENIKE.

T h e larva is s imi lar except in size to t h a t of P. coccinea KOCH. P . 4 bears two br is t les app rox ima te ly half the length of the long br is t le on P . 3. E. I cornes to a s h a r p po in t r each ing pas t the po in t br is t le of inse r t ion of the pa lp i .

T h e s u t u r e be tween E. II & E. III is a lmos t equ id i s t an t be tween the rea r edge of E. I and the br is t le E p . 4. T h e excre to ry p la te

( 5 ) LARVAE OF HYDRACARINA 235

F I G . 3 : Aturus scaber KRAMER, larva , palpi . — F I G . 4 : id°, excretory field.

is t r i a n g u l a r w i th well r o u n d e d angles . I t is r a t h e r longer t h a n b road . T h e r ea r end of E . III déviâtes r o u n d t h e br i s t le Ex. 1.

M e a s u r e m e n t s (ju) : 1 2 3 4

Leng th . B r e a d t h

395 395 425 400 220 195 220 220

2 3 6 R. K. H. JONES ( 6 )

Capi tu lum 1 2 0 1 2 0 1 3 0 1 2 0

Ex. 4 1 5 5 1 3 0 1 5 5 1 5 5

1 st leg 2 9 0 2 9 0 3 1 0 3 1 0

2 nd leg 3 4 0 3 2 5 3 4 0 3 2 5

3 rd leg 3 2 5 3 6 0 3 6 0 3 5 0

Excre to ry p la te :

Leng th 4 5 4 5 5 0 4 5

Bread th 4 0 4 0 4 5 4 0

F I G . 5 : Piona uncata K. , larva, ventra l surface . — F I G . 6 : id°, excretory fleld.

2 . 3 . — Protzia eximia PROTZ.

On the an t e r i o r pa r t of the d o r s u m is a not very well delined p la te bear ing the méd ian eye and th ree pa i r s of long br is t les , the middle pa i r of which are pec t ina te and the o the r two pa i r s s imple . T h e pa i red eyes a re large and close to the edge of the p la te . T h e r ea r eye of each p a i r is no t d is t inc t ly p igmented .

Db. 1 to 8 a re ail pec t ina te . Db. 8 a re on rounded p r o t r u b e r a n c e s at t he ex t rême pos te r io r end of the larva .

T h e legs a re compara t ive ly long and bear large n u m b e r s of long pec t ina te br is t les . T h e r ea r side of segment 2 of the second leg and the p rox ima l dorsa l p a r t of segment 1 of the th i rd leg

(7) LARVAE OF HYDRACARINA 237

each bea r a s imple br is t le . The c laws consis t of a very large m é d i a n c law wi th two smal le r c laws, one on e i ther side.

T h e palp i bea r a s t rong curved spine on P . 2 and a br is t led p r o t r u b e r a n c e on P . 3 . T h e r e a re only 4 visible segments .

M e a s u r e m e n t s (p.) : 1 2 3 4

250 240 290 290 B r e a d t h 130 120 120 130

50 70 50 50 Pa lp i 90 100 90 85 1 st leg 310 380 360 310 2 nd leg 290 290 310 290 3 rd leg 325 375 340 325

F I G . 7 : Protzia eximia PROTZ, larva , dorsal surface .

238 R. K. H . JONES ( 8 )

3. — L I S T O F K N O W N H O S T S O F W A T E R - M I T E S .

In the table following t h e ear l ies t k n o w n a u t h o r is given in each case. W h e r e subséquen t a u t h o r s have given différent hos t s thèse a re also l isted.

T h e n a m e s of hos t s a re given exact ly as in the p a p e r s cited and no a t t e m p t h a s been m a d e to b r ing t h e m u p to da te , bu t in the case of the mi tes themselves the s y n o n y m s and classification have been b r o u g h t in to line w i t h V I E T S [ 1 9 5 6 ] .

I have been able to find n o in fo rmat ion wha t soeve r on hos t s of the super- famil ies Mideopsae VIETS or Krendowsk iae V I E T S .

M i t e C l a s s i f i c a t i o n HOST. [ H o s t c lass i f i cat ion] A U T H O R

1 . S u p e r - f a m i l y H y d r o -v o l z i a e V I E T S

H y d r o v o l z i i d a e T H O R Hydrouolzia T H O R H. gerhardi M I T C H E L L MICROVELIA AMERICANA

U H L E R [Ins . H e m . ] M I T C H E L L 1 9 5 4

Z. S u p e r - f a m i l y H y -d r a c h n a e Part .

H y d r a c h n i d a e L E E C H Hydrachna M U L L E R H. cruenta M U L L E R H. globosa D E G E E R

H. magniscutata MARSCHALL

H. maramauensis STOUT

H. processifera KOENIKE

H. uniscutata T H O R H. leegei KOENIKE

CORIXINAE [ Ins . H e m . ] Dorsa l on DYTISCS. [ Ins . Col .] NEPA; RANATRA [ Ins . H e m . ] DYTISCIDAE [ Ins . Col .] CYMATIA; NOTONECTA

[ Ins . H e m . ]

NOTONECTA UNDULATA; RANATRA; ARCTOCORIXA

[ Ins . H e m . ] ANISOPS WAKEFIELDI ; ARCTOCORIXA ARGUTA

[ Ins . H e m . ] DYTISCUS MARGINALIS L.

[ Ins . Col . ] DYTISCS [ Ins . Col . ] NOTONECTA GLAUCA L.

[ Ins . H e m . ]

SPARING 1 9 5 9

W E S E N B E R G - L U N D 1 9 1 8

W E S E N B E R G - L U N D 1 9 1 8

V I E T S 1 9 3 6 d

V I E T S 1 9 3 6 d

CROWELL 1 9 6 0

STOUT 1 9 5 3 b

W E S E N B E R G - L U N D 1 9 1 8

W E S E N B E R G - L U N D 1 9 1 8

JONES 1 9 6 5

. S u p e r - f a m i l y L i m n o c h a r a e V I E T S

L l m n o c h a r i d a e G R U B E : Limnochares LATREII .LE Limnochares aquatica L. GERRIDS [ Ins . H e m . ]

VELIIDS & HYDROMETRIDS [ Ins . H e m . ]

AQUARIUS PALLUDUM & LIMNOPHORVS RUFOSCU-TELLATUS [ Ins . H e m . ]

ZYGOPTERAN IMAGINES [ Ins . Odo.]

D U G E S 1 8 3 4 b

SPARING 1 9 5 9

IMAMURA 1 9 5 4 b

KILLINGTON & B A T H E 1 9 4 7

(9) LARVAE OF HYDRACARINA 239

E y l a i d a e L B B C H : Eylais LATREILLE E. bisinuosa KOENIKE

E, extendens M Û L L E R E. discreta KOENIKE E. hamata KOENIKE

E. infundibulifera KOENIKE

do meridionalis T H O N E. mutila KOENIKE

E. setosa KOENIKE

E. waikewai STOUT

P r o t z i i d a e V I E T S : Protzia P I E R S I G P. eximia PROTZ

Partnunia P I E R S I G P. steinmanni W A L T E R P. achidai IMAMURA

CYMATIA COLEOPTRATA F A B . [ Ins . H e m . ]

CORIXINAE [ Ins . H e m . ] CORIXINAE [ Ins . H e m . ] CYMATIA [ Ins . H e m . ] DYTISCUS; COLEMBETES;

GYR1NUS; HYDROPHILUS; GR A PHODER ES ; RANTUS; ILYBIUS; HYPHYDRUS.

[ Ins . Col . ] GILLS OF FISHES [Cho. P i s . ] DYTISCS & GYRINIDS

[ Ins . H e m . ] CORIXA [ Ins . H e m . ] DYTISCS & GYRINIDS

[ Ins . Col . ] DYTISCS & GYRINIDS

[ Ins . Col . ] ANISOPS WAKEFIELDI

[ Ins . H e m . ]

JONES 1 9 6 5

SPARING 1 9 5 9

SPARING 1 9 5 9

V I E T S 1 9 3 6 d

V I E T S 1936 d

V I E T S 1936 d

PIATAKOV 1915

BOTTGER 1962

PIATAKOV 1915

PIATAKOV 1915

STOUT 1953 a

WEIDEMANIA BISTIGMATA C U R T I S [ Ins . Dip . ] JONES 1965

UNIDENTIFIED CADDIS FLY [ Ins . Tri . ] JONES 1965

PLECOPTERA [ Ins . P ie . ] 6 Spp. of PLECOPTERA IMA­

GINES [ Ins . P ie . ]

MUNCHBERG 1954

IMAMURA 1954 b

4. S u p e r - f a n a l l y H y d r y -p h a n t a e V I E T S

T h y a s i d a e T H O R : Thyasides LUNDBLAD T. dentata T H O R Panisopsis V I E T S P. curvifrons W A L T E R

P. n. spp ( u n n a m e d )

Thyopsis P I E R S I G T. cancellata PROTZ Thyas KOCH T. barbigera V I E T S T. spp. P r o b a b l y T.

pachystoma P I E R S I G T. stolli KOENIKE

Trichothyas V I E T S T. muscicola M I T C H E L L

Panisus KOENIKE. P. michaeli KOENIKE

H y d r y p h a n t i d a e T H O R : Hydryphantes KOCH H. ruber D E GEER

AEDES LUTESCENS [ Ins . Dip . ] LUNDBLAD 1927 c

DIPTERAN IMAGINES [ Ins . Dip . ]

APTEROUS STONEFLIES [ Ins . P i e . ]

Var ious TIPULIDS [ Ins . Dip . ]

TIPULIDS [ Ins . D ip . ]

TIPULIDS [ Ins . Dip . ]

1. AEDES VEXANS & AEDES spp. [ Ins . Dip . ]

2. Hatched as n y m p h s

LIMONIA HUMIDICOLA O. S. [ Ins . D ip . ]

GYROCAMPA UBIGINOSA H A L . [ Ins . H y m . ]

CVLEX [ Ins . Dip . ]

W A L T E R 1915 a

W O M E R S L E Y 1953

MUNCHBERG 1935 b

MUNCHBERG 1935 b

MUNCHBERG 1935 b

C R O W E L L 1960

C R O W E L L 1960

M I T C H E L L 1953

LUNDBLAD 1927 c

V I E T S 1936 d

240 R . K . H . J O N E S (10)

PALUDINA [Mol. Gas . ] H. dispar VON S C H A U B EPHYDRINAE [ Ins . Dip . ] Georgella KOENIKE G. helvatica HALLER ISCHNURA; AGRION

[ Ins . Odo.] H y d r o d r o m i d a e V I E T S Hydrodroma KOCH H. despiciens M U L L E R CHIRONOMIDS; CORETHRA

[ Ins . D ip . ]

T H O R 1 8 9 9

SOAR 1 9 0 6

MÙNCHBERG 1 9 3 5

W E S E N B E R G - L U N D 1 9 1 8

5 . S u p e r - f a m i l y L e b e r -t i a e V I E T S

Sperchon idae T H O R : Sperchon

S. brevirostris KOENIKE

S . denticulatus KOENIKE S . setiger T H O R

5 . squamosus KRAMER

S . glandulosus KOENIKE

S . longissimus V I E T S

S . jasperensis M A R S H A L L S . spp. Lebert i idae T H O R : Lebertia NBUMANN L. stigmatifera T H O R

L. glabra T H O R Torrent i co l idae

P I E R S I G :

Torrenticola P I E R S I G T. spongicola V I E T S

ORTHOCLADIUS SORDIDEL-LUS ZETTERSTADT [ Ins . D ip . ]

CHIRONOMIDS [ Ins . Dip . ] P u p a e of MELUSINA ORNATA

[ Ins . D ip . ] ORTHOCLADIUS SORDIDEL-

LUS [ Ins . D ip . ] 4 spp. CHIRONOMIDS

[ Ins . D ip . ] ORTHOCLADIUS FLAVIDULA

E D W A R D S [Ins . Dip . ]

SIMULIUM spp. [ Ins . D ip . ] CHIRONOMIDS [ Ins . D ip . ]

Direct d e v e l o p m e n t to n y m p h n o n e

2 spp CHIRONOMIDS [ Ins . Dip . ]

SPONGES [ P o r i f e r a ]

THIENNEMANN 1 9 0 8

SPARING 1 9 5 9

BESSELING 1 9 3 9

V I E T S 1 9 3 6 d

EFFORD 1 9 6 0

EFFORD 1 9 6 0

D A V I E S 1 9 5 9

JONES 1 9 6 5

LUNDBLAD 1 9 2 4

EFFORD 1 9 6 0

6 . S u p e r - f a m i l y P i o n a e V I E T S

P o n t a r a c h n i d a e T H O R : Pontarachna P H I L I P P I P. punctulum P H I L I P P I OSTREA [Mol. Lam.] ANDRÉ & LAMY 1 9 3 0

L i m n e s i i d a e T H O R : Limnesia KOCH L. connata KOENIKE Hatched as n y m p h P I E R S I G 1 9 0 0 L. undulata M U L L E R Hatched as n y m p h P I E R S I G 1 9 0 0

TANYTARSUS DECORUS JOHANNSEN [ Ins . Dip . ] CROWELL 1 9 6 0

L. fulgida KOCH CHIRONOMIDS [ Ins . Dip . ] SPARING 1 9 5 9 WATER-BEETLE LARVAE

[ Ins . Col . ] W E S E N B E R G - L U N D 1 9 1 8

CHIRONOMIDS [ Ins . Dip . ] SPARING 1 9 5 9 L. maculata M U L L E R CHIRONOMIDS [ Ins . D ip . ] LUNDBLAD 1 9 2 6 c H y g r o b a t i d a e KOCH : Hygrobates KOCH H. foreli LEBBRT CHIRONOMIDS [ Ins . D ip . ] LUNDBLAD 1 9 2 7 c H. tongipalpis HERMANN CORETHRA; CHIRONOMIDS

[ Ins . Dip . ] MÙNCHBERG 1 9 3 5 b

(•n : LARVAE OF HYDRACARINA 241

Atractides KOCH A. nodipalpis T H O R A. spinipes KOCH

U n i o n i c o l i d a e OUDEMANS :

Vrtionicola HALDEMAN V. aculeata KOENIKE

Aduts & n y m p h s

Larvae U, abnorpipes WOLCOTT

U. arcuata WOLCOTT

U. bonzi CLAPARÈDE

U. cirrosa KOENIKE

U. ampullariae KOENIKE U. crassipes M U L L E R U. diversipes V I E T S U. flssipes KOENIKE

U. formosa DANA & W H E L P L E Y

U. fossulata KOENIKE

U. intermedia KOENIKE Aduts & n y m p h s

Larvae

U. japonensis V I E T S

U. parasitica U C H I D A & IMAMURA

U. perforatus KOENIKE

U. procurvipes KOENIKE

U. rugosa KOENIKE

U. uchidai IMAMURA U. serrata WOLCOTT

[7. ypsilophora BONZ

Polyatacides LUNDBLAD P. prominens KOENIKE Atacella LUNDBLAD. A. schubarti V I E T S

A. clathrata LUNDBLAD Najadicola P I E R S I G N. ingénus KOENIKE

Neumania LEBERT N. deltoides PIERSIG

MIDGE IMAGINES [ Ins . Dip . V a r i o u s CHIRONOMIDS

[ Ins . Dip .

Not paras i t i c b u t res t ing stages found in MUSSELS

CAMPELOMA LECISIUM S A Y [Mol. Lam. ]

ANODONTA ANATINA L. [Mol. Lam. ]

V a r i o u s CHIRONOMIDS LAMPSILLIS SI LI QUOI DES

[Mol. Lam. ] LAMPSILLIS SILIQUOIDES

[Mol. Lam.] UNIO DOUGLASSIAE NIP-

PONENSIS [Mol. Lam. ] UNIO NEPEANENSIS

[Mol. Lam. ] AMPULLARIA [Mol. Gas.] F. W. SPONGES [ P o r i f e r a ] NAÏADES [Mol. Lam. ] ANODONTA LATIMARGINATA

[Mol. Lam. 4 ANODONTA SPECIES

[Mol. Lam. LAMPSILLIS SILIQUOIDES

[Mol. Lam.

ANODONTA ANATINA L. [Mol. Lam.

V a r i o u s CHIRONOMIDS [ Ins . Dip.

VIVIPAROUS MALLEATUS [Mol. Gas.

ANODONTA BERINGUANA [Mol. L a m .

ANODONTA LATIMARGINATA [Mol. Lam.

ANODONTA GIGANTEA [Mol. L a m .

ANODONTA LATIMARGINATA [Mol. Lam.

F. W. MUSSELS [Mol. Lam. ] LAMPSILLIS SILIQUOIDES

[Mol. Lam. ANODONTA CYGNEA L.

[Mol. Lam.

F. W. MUSSELS [Mol. Lam.]

DIPLODON DELONTUS EX­PANSUS KUSTER [Mol. Lam. ]

F. W. MUSSELS [Mol. L a m . ]

3 gênera of MUSSELS [Mol. L a m .

CHIRONOMIDS [ Ins . Dip . ]

IMAMURA 1 9 5 4 b

JONES 1 9 6 5

M I T C H E L L 1 9 5 5

CHANDLER 1 9 3 4

JONES 1 9 6 5

JONES 1 9 6 5 b

M I T C H E L L 1 9 5 5

M I T C H E L L 1 9 5 5

IMAMURA 1 9 5 3

KOENIKE

ANDRÉ & LAMY 1 9 3 0

ARNDT & V I E T S 1 9 3 8

V I E T S 1 9 2 6

ANDRÉ & LAMY 1 9 3 0

M I T C H E L L 1 9 5 7

M I T C H E L L 1 9 5 5

MUTCHELL 1 9 5 5

JONES 1 9 6 5 b

IMAMURA 1 9 5 3

IMAMURA 1 9 5 4 b

ANDRÉ & LAMY 1 9 3 0

ANDRÉ & LAMY 1 9 3 0

ANDRÉ & LAMY 1 9 3 0

IMAMURA 1 9 5 3

M I T C H E L L 1 9 5 5

M I T C H E L L 1 9 5 5

LUNDBLAD 1 9 3 7

V I E T S 1 9 5 4

LUNDBLAD 1 9 3 7

M I T C H E L L 1 9 5 5

SPARING 1 9 5 9

2

2 4 2 R . K . H . J O N E S (12)

N. uchidai I M A M U R A

Huitfeldtia T H O R H. rectipes F e l t r i i d a e T H O R : Feltria K O E N I K E

F. setigera ( ? ) K O E N I K E

F. romijni B E S S E L I N G

F. minuta K O E N I K E

P i o n i d a e T H O R :

Typhis K O C H T. ornatus K O C H

T. spp.

Pionopsis P I E R S I G

P. lutescens H E R M A N N

Pionocercus P I E R S I G

Pionocercus P I E R S I G .

P. leuckarti P I E R S I G

Piona K O C H P. carnea K O C H

P. coccinea K O C H P. conglobata K O C H

P. disparalis K O E N I K E

P. linguaplax C R O W E L L

P. nodata M U L L E R

P. reighardi W O L C O T T

P. rotunda K R A M E R

P. uncata uncata K O E N I K E

P. uncata exigua V I E T S

P. variabilis K O C H Foreil H A L L E R

F. spp.

7. S u p e r - f a m i l y A x o -n o p s a e

A x o n o p s i d a e V I E T S :

Brachypoda L E R E R T

B. versicolor M I Ù L L E R

Ljania T H O R L. bipapillata T H O R Parasitalbia V I E T S

P. sumatrensis V I E T S

8 . S u p e r - f a m i l y A r r e -n u r a e

Arrenuridae T H O R : Arrenurus D U G È S

S u b - g e n u s Arrenurus str. D U G È S

MIDGES [ Ins . D i p . ]

CHIRONOMIDS [ Ins . D ip . ]

CHIRONOMIDS [ Ins . D ip . ] TANYTARSUS FLAYIDULUS

E D W A R D S [ Ins . D ip . ]

Several spp. CHIRONOMIDS

CHIRONOMUS THUMMI PV-PAE [ Ins . D ip . ]

Var ious CHIRONOMUS IMA­GINES [ Ins . D ip . ]

CHIRONOMIDS [ Ins . Dip . ]

I M A M U R A 1 9 5 4 b

M Ù N C H B E R G 1 9 3 5

V I T Z H U M 1 9 4 1

E F F O R D 1 9 6 0

E F F O R D 1 9 6 0

U C H I D A 1 9 3 2

J O N E S 1 9 6 5

S P A R I N G 1 9 5 9

Direct d e v e l o p m e n t to n y m p h L U N D B L A D 1 9 2 7 c

CHIRONOMUS THUMMI PU-PAE [ Ins . D ip . ]

CHIRONOMIDS [ Ins . D ip . ] CHIRONOMIDS [ Ins . D i p . ] CHIRONOMUS DORSALUS

M E I G E N [ Ins . Dip . ]

CRICOTOPUS snp. [ Ins . D ip . ] POLYPEDILUM spp. [ Ins . Dip . ] Direct d e v e l o p m e n t to n y m p h . Larvae large t y p e to n y m p h . Larvae s m a l l t y p e UNKNOWN

HOST TENDIPES DECORUS

J O H A N N S E N [ Ins . D ip . ]

Direct d e v e l o p m e n t to n y m p h CHIRONOMUS PLUMOSUS L.

OTHER CHIRONOMIDS [ Ins . D ip . ]

Direct d e v e l o p m e n t to n y m p h CHIRONOMIDS [ Ins . D ip . ]

U C H I D A 1 9 3 2

S P A R I N G 1 9 5 9

S P A R I N G 1 9 5 9

J O N E S 1 9 6 5

M O T A S 1 9 2 8

C R O W E L L 1 9 5 3

P I E R S I G 1 9 0 0

B Ô T T G E R 1 9 6 2

B Ô T T G E R 1 9 6 2

C R O W E L L 1 9 6 0

P I E R S I G 1 9 0 0

J O N E S 1 9 6 5

S T O U T 1 9 5 3

J O N E S 1 9 6 5

N y m p h s h a t c h e d f r o m eggs. C O O K 1 9 5 6

Hatches as n y m p h P I E R S I G 1 9 0 0

4 spec ies CHIRONOMIDS E F F O R D 1 9 6 0

As a d u l t s on EPHEMERW LARVAE [ Ins . Eph . ] V I E T S 1 9 3 5

(13) LARVAE OF HYDRACARINA 243

A. abbreviator B E R L E S B LIBELLULIDAE [ Ins . Odo.] A. affinis K ΠN I K E LIBELLULIDAE [ Ins . Odo.] A. agrionicolus U C H I D A . AGRIONIDAE [ Ins . Odo.] A. Americanus major ISCHNURA VERTICALIS S A Y

M A R S H A L L [ Ins . Odo.]

A. angelieri Cass. - ZYGOPTERA [ Ins . Odo.] MÉJEAN

A. batillifer K ΠN I K E LIBELLULIDAE [ Ins . Odo.] .4. bicuspidator B E R L E S B LIBELLULIDAE [ Ins . Odo.] .4. bleptopetiolatus EPICIRDULIA PRINCEPS

COOK HAGEN [Ins . Odo.]

A. bruzelii K ΠN I K E LIBELLULIDAE [ Ins . Odo.] A. claviger K ΠN I K E LIBELLULIDAE [ Ins . Odo.] A. compactais ENALLAGMA EBRIUM HAGEN

M A R S H A L L [ Ins . Odo.]

A. compactus P I E R S I G LIBELLULIDAE [ Ins . Odo.] .4. crenatus K ΠN I K E LIBELLULIDAE [ Ins . Odo.] .4. crassicaudatus CORETHRA CRISTALLIN A

KRAMER D E GEER [Ins . Dip . ]

ANOPHELES MACULIPENNIS MBIGEN [ Ins . Dip . ]

A. cuspidator M U L L E R LIBELLULIDAE [ Ins . Odo.] A. cuspidifer P I E R S I G LIBELLULIDAE [ Ins . Odo.] A. daisetsuenses AESCHNA NIGROFLAVA

IMAMURA ENALLAGMA DESERTI [ Ins . Odo.]

A. dimontifer ZYGOPTERA [ Ins . Odo.] MÙNCHBERG

A. fimbriatus K ΠN I K E ANOPHELES MACULIPENNIS MlEIGEN

THEOBALDIA ANNULAT A [ Ins . Dip . ]

A. fissicornis M A R S H A L L LIBELLULIDAE [ Ins . Odo.] A. furcillatus V I E T S ZYGOPTERA [ Ins . Odo.] A. lattis BARROIS & ANOPHELES MACULIPENNIS

MONIEZ MEIGEN [ Ins . D ip . ]

.4. leuckarti P I E R S I G LIBELLULIDAE [ Ins . Odo.] A. maculator M U L L E R LIBELLULIDAE [ Ins . Odo.]

ZYGOPTERA [ Ins . Odo.] A. munchbergi K . O.

V I E T S & CASSAGNE-

M É J E A N ZYGOPTERA [ Ins . Odo.] A. neumani P I E R S I G LIBELLULIDAE [ Ins . Odo.] A. octotamuli

MÙNCHBERG ZYGOPTERA [ Ins . Odo.] A. ornatus GEORGE ZYGOPTERA [ Ins . Odo.] A. planus MIARSHALI. ANISOPTERA & ZYGOPTERA

[ Ins . Odo.] .4. platg-rotundo-cuspi- ANISOPTERA & ZYGOPTERA

dator MÙNCHBERG [ Ins . Odo.] A. papillator M U L L E R LIBELLULIDAE [ Ins . Odo.] A. pollictus M A R S H A L L 4 LESTES spec ies [ Ins . Odo.] A. pustulator M U L L E R LIBELLULIDAE [ Ins . Odo.] A. radiatus P I E R S I G LIBELLULIDAE [ Ins . Odo.] A. robustus K Œ N I K E LIBELLULIDAE [ Ins . Odo.] .4. rodrigensis LUNDBLAD ZYGOPTERA [ Ins . Odo.] .4. tetracyphus P I E R S I G LIBELLULIDAE [ Ins . Odo.] .4. tricuspidator M U L L E R LIBELLULIDAE [ Ins . Odo.] .4. virens NBUMAN LIBELLULIDAE [ Ins . Odo.] .4. wallensis COOK LIBELLULIDAE [ Ins . Odo.]" Sub-genus Megaluraca-

rus V I E T S

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

IMAMURA 1958

MÙNCHBERG 1951

C A S S . - M É J E A N 1966

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

M I T C H E L L 1959

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

M I T C H E L L 1959

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

MONCHBERG 1935 a

MÙNCHBERG 1958

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

IMAMURA 1951 a

MÙNCHBERG 1958 b

MÙNCHBERG 1936 e

M I T C H E L L 1959

C A S S . - M É J E A N

MÙNCHBERG 1938

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

JONES 1965

C A S S . - M É J E A N 1966

MÙNCHBERG 1935 d

M'ÙNCHBERG 1958 b

KILLINGTON & B A T H E 1947 a

MÙNCHBERG 1952

MÙNCHBERG 1952

MÙNCHBERG 1935 d

M I T C H E L L 1959

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

C A S S . - M É J E A N 1966

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

MÙNCHBERG 1935 d

M I T C H E L L 1959

244 R. K. H . JONES (14)

A. buccinator M U L L E R

A. einbergeri MUNCHBERG

A. globator M U L L E R

A. medio-rotundatus T H O R

A. odonatophilus MUNCHBERG

S u b - g e n u s Truncaturus T H O R

A. knauthi K ΠN I K E . 4 . nodosus K ΠN I K E A. stecki K ΠN I K E A. truncatellus

M U L L E R

S u b - g e n u s Micruracanis V I E T S

. 4 . bisulcicodulus P I E R S I G

A. integrator M U L L E R A. pugionifer K ΠN I K E . 4 . sinuator M U L L E R

Sub-genus uncer ta in , probab ly Arrenurus s. str.

A. imperator M U L L E R A. retroduobus

MUNCHBERG

S u b - g e n u s u n d e t e r m i -ned

.4. species A. species 1

. 4 . Species 2

Various MOSQUITOES [ Ins . Odo.]

UBELLULIDAE [ Ins . Odo.] 2 Var ious MOSQUITOES

[ Ins . Dip. ] CULEX PIPIENS LARVAE

[ Ins . Dip . ] DYTISCUS & CYRINUS

LARVAE [ Ins . Col.] ANOPHELES CLAVIGER

MEIGEN [Ins . Dip . ]

UBELLULIDAE [ Ins . Odo.]

MOSQUITOES [ Ins . Dip . ] MOSQUITOES [ Ins . Dip . ] MOSQUITOES [ Ins . Dip . ]

MOSQUITOES [ Ins . Odo.]

MOSQUITOES [ Ins . Dip . ] MOSQUITOES [ Ins . D ip . ] MOSQUITOES [ Ins . D ip . ] TANNYPUS PUNCTIPENNIS

MEIGEN [Ins . D ip . ]

UBELLULIDAE [ Ins . Odo.]

UBELLULIDAE [ Ins . Odo.]

Young fish [Cho. P i s . ] DIXA AESTIVALIS MEIGEN

[ Ins . Dip . ] ANATOPYNIA VARIA JOHNSTON

[ Ins . Dip. ]

M U N C H B E R G 1 9 3 5 b

M U N C H B E R G 1 9 6 0

M U N C H B E R G 1 9 3 6 e

BÔTTGBR 1 9 6 2

JONES 1 9 6 5

MUNCHBERG 1 9 5 8

MUNCHBERG 1 9 3 6 e

MUNCHBERG 1 9 3 6 e

MUNCHBERG 1 9 3 6 e

MIÙNCHBERG 1 9 3 7 b

MUNCHBERG 1 9 3 7 b

MUNCHBERG 1 9 3 8 a

MUNCHBERG 1 9 3 6 e

MUNCHBERG 1 9 3 6 e

MUNCHBERG 1 9 6 0

MUNCHBERG 1 9 5 9

SOAR * WlLLIAMSON 1 9 2 5

JONES 1 9 6 5

JONES 1 9 6 5

SUMMARY

The larvae of thèse three mites are described for the first time.

1. — Aturus scaber Kramer.

Females of this species lay one, very large egg at a time, suggesting a suppression of the larval stage and direct development to nymph. This expectation has been refuted and the larvae bred in the laboratory. Further information on the life-history is lacldng but a parasatic stage is indicated.

2 . — Piona uncata KOENIKE. The complète life-history of this mite is described. In the spécimens

studied the main host was Chironomus plumosus L. "Work by STOUT shows that this species has an alternative direct development. The larvae are similar except in size to those of P. coccinea KOCH.

(15) LARVAE OF HYDRACARINA 245

3. — Protzia eximia Protz.

Eggs are found throughout the winter and laboratory tests show that hatching is dépendant on température. The larvae resemble those of Panisopsis species, and run actively on the water film. They are able to jump 2 cm from the surface and use this ability to gain a place on their hosts, an unidentifled Trichopteran and the fly, Weidemannia bistigmata CURTIS. There are probably two générations a year.

An up-to-date table gives hosts for 150 species and références for this information.

D E S C R I P T I O N D E S L A R V E S O'ATURUS SCABER KRAMER, PROTZIA EXIMIA PROTZ E T PIONA UNCATA K ΠN I K E

E T N O T E S S U R L E C Y C L E B I O L O G I Q U E D E S D E U X D E B N I È B E S E S P È C E S .

[Hydracarina]

Les larves de ces trois Hydracariens sont décrites pour la pre­mière fois.

1. — Aturus scaber KRAMER.

Les femelles de cette espèce pondent un très gros œuf unique, fait qui avait suggéré l'absence du stade larvaire et un développement don­nant directement une nymphe. Cette supposition s'est avérée inexacte et les larves ont été élevées au laboratoire. On manque encore d'autres données sur le cycle biologique, mais il semble exister un stade parasite.

2. — Piona uncata KŒNIKE.

Le cycle biologique complet de cette espèce est décrit. Chez les spé­cimens étudiés, l'hôte principal est Chironomus plumosus L. Un travail de STOUT montre que cette espèce peut également présenter un déve­loppement direct. A l'exception de leur taille, les larves ressemblent à celles de P. coccinea KOCH.

3. — Protzia eximia PROTZ. On rencontre des oeufs de cette espèce pendant tout l'hiver et des

expériences réalisées au laboratoire montrent que Péclosion dépend de la température. Les larves ressemblent à celles des espèces de Panisopsis et courent activement à la surface de l'eau. Elles sont capables de sauter à 2 cm de la surface et utilisent cette faculté pour parvenir sur leurs hôtes, un Trichoptère non identifié et le Diptère Weidemannia bistigmata CURTIS. Il existe probablement deux générations par an.

Une table mise à jour indique les hôtes de 150 espèces et les références bibliographiques de ces données.

B E S C H B E I B U N G D E E L A E V E N V O N ATURUS SCABER KRAMKR, PROTZIA EXIMIA PROTZ U N D PIONA UNCATA K ΠN I K E

U N D A N G A B E N Û B E E D E N E N T W I C K L U N G S Z Y K L U S D E E B E I D E N L E T Z T E N A B T E N .

[Hydracarina]

Die Larven dieser drei Hydracarinen werden zum erstenmal beschrieben.

1. — Aturus scaber KRAMER. Bci dieser Ar t legen die Weibchen ein

246 R. K. H . JONES ( 1 6 )

einziges grosses Ei, wodurch man auf das Fehlen des Larvenstadiums und auf eine direkte Entwicklung zur Puppe geschlossen hatte.

Dièse Annahme bezeigte sich als ungenau, und die Larven wurden im Laboratorium aufgezogen. Weitere Angaben iiber den Entwicklungs-zyklus fehlen noch, aber es existiert auscheinend ein Parasitenstadium.

2. — Piona uncata KŒNIKE . Der gesamte Entwicklungsgang dieser Art wird beschrieben. Bei den bearbeiteten Exemplaren ist der hauptsâchlichste Wirt Chironomus plumosus L.

Eine Verôffentlichung von STOUT zeigt, dass es bei dieser Art ebenfalls eine direkte Entwicklung geben kann. Wenn man von ihrer grosse absieht, âhneln dièse Larven denen von P. coccinea KOCH.

3. — Protzia eximia PROTZ. Wahrend des ganzen Winters kann man Eier von dieser Art flnden und die im Labor durchgefùhrten Versuche zeigen, dass das Schliipfen von der Temperatur abhângt.

Die Larven âhneln denen der verschiedenen Arten von Panisopsis und bewegen sich aktiv an der Wasser-oberflâche. Sie kônnen Springe bis zu 2 cm ùber dem Wasserspiegel machen und benûtzen dièse Fâhig-keit, um auf ihre "Wirte, eine nicht bestimmte Kôcherfliege und den Zweifliigler Weidemannia bistigmata CURTIS ZU gelangen. Wahrschein-lich gibt es jahrlich zwei generationen.

Eine vervollstândigte Tafel zeigt die Wirte von 150 Arten sowie aile diesbeziiglichen Literaturnachweise auf.

B I B L I O G R A P H Y

For convenience, ail références before 1955 are referred to KARL VIETS :

VIETS ( K . ) . 1955. — Die Milben des Susswassers und des Meeres. Teil I .

Bibliographie. Gustav Fischer. Jena.

Other références :

BOTTGER ( K . ) . 1962. — Zur Biologie und Ethnologie der einheimischen Wassermilben Arrenurus (M) globator MULLER, Piona nodala nodata MULLER und Eylais infundibuli'era meridionalis T H O N . Zool. Jb. SysL, 8 9 : 501-584.

— 1965. — Das parasitare Larvenstadium von A. valdiviensis K . O. VIETS. Z. Morph. Okol., 5 5 ; 383-409.

CASSAGNE-MÉJEAN (F.). 1966. — Contribution à l'étude des Arrenuridae de France. Acarologia, 8 .

CooK ( D . B.). 1955. — Preliminary Studies of the Hydracarina of Michi-gan. The sub-familv Foreliinae VIETS. Ann. ent. Soc. Amer., 4 8

(4) : 299-307. CROWELL (B. M.). 1960. — Taxonomy, Distribution and Development

stages of Ohio Water Mites. Bull. Ohio. Biol. Soc, 1 (2). DAVIES (M. D . ) . 1959. — The Parasitism of Black Flies (Diptera, Simu-

liidae) by larval water mites mainly of the genus Sperchon. Canad. J. Zool., 3 7 : 353-369.

EFFORD (il. E.). 1960. — A Population Study on Water-Mites. Unpublished Thesis for Degree of Ph. D. at Ox'ord University.

IMAMURA (T.). 1958. — Water Mites of Marsh Hinuma. Bull. Fac. Lib. Arts. Ibaraki Univ., 8 .

(17) LARVAE OF HYDRACARINA 247

JONES ( R . K. H.). 1965 a. — Ecology of the Hydracarina. Unpublished Thesis for the Degree of Ph. D. at Leicester University.

JONES (R. K. H.). 1965 b. — Parasitism by Larvae of Unionicola inter-media Koenike and another Unionicola Sp. (Acarina, Pionae on Chironomids. Nature, Lond., 207 (4994) : 317-318.

MITCHELL ( R . D.). 1955. — Anatomy, Life-History and Evolution of the Mites Parasitising Fresh-Water Mussels. Mise. Publ. Mus. Zool. Univ. Mich. 8 9 .

MITCHELL ( R . D.). 1957. — On the Mites Parasitising Anodonta (Unioni-dae : Mollusca). J. Parasit., 4 3 (1) : 101-104.

MITCHELL ( R . D.). 1959. — Life-histories and Larval Behaviour of Arrenurid Water-mites. J. N. Y. Ent. Soc, 67 (1).

MÙNCHBERG (P.). 1958 a. — Eine unbekannte Megaluracarus-Arï (Hydra-chnellae, Acari) aus Belgisch Kongo mit libellenparasitischer Larvenphase. Zool. Anz., 160 : 47.

MÙNCHBERG (P.). 1958 b. — Erster Beitrag iiber die an Afrikanischen Odonaten Parasitierenden Arrenurus - larven. (Hydrachnellae, Acari). Z. Parasitenk. 18 ; 243-256.

MÙNCHBERG (P.). 1959. — Dritte Beitrag iiber die an afrikanischen Libellen Schmarotzenden Arrenurus - Larven. (Hydrachnellae, Acari). Arch. Hydrobiol., 55 : 264-275.

MÙNCHBERG (P.). 1960. — Erste Mitteilung iiber die an brasilianischen Odonaten schmarotzenden Arrenurus - Larven (Hydrachnellae, Acari). Arch. Hydrobiol., 5 6 (4) : 425-440.

SPARING (L). 1959. — Die Larven der Hydrachnellae, ihre parasitische Entwicklung und ihre Systematik. Parasitologische Schriftenreihe, 10.

VIETS (K.). 1956. — Die Milben des Siisswassers und des Meeres. Teil H et III. Katalog und Nomenklator. Gustav Fischer. Jena.

(Jordanhill Collège, Glasgow, Scotland).