Mook, Jacob Hendrik

University of Groningen

   University of GroningenHabitat selection by Lipara lucens Mg. (Diptera, Chloropidae) and its survival valueMook, Jacob HendrikIMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite fromit. Please check the document version below.Document VersionPublisher's PDF, also known as Version of recordPublication date:1967Link to publication in University of Groningen/UMCG research databaseCitation for published version (APA):Mook, J. H. (1967). Habitat selection by Lipara lucens Mg. (Diptera, Chloropidae) and its survival value.Groningen: Noordhoff Uitgevers.CopyrightOther than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of theauthor(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons).Take-down policyIf you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediatelyand investigate your claim.Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons thenumber of authors shown on this cover page is limited to 10 maximum.Download date: 12-11-2019545 HABITAT SELECTION BY LIPARA LUCENS MG. This tendency to cluster, however, will be counteracted to some extent by the randomness in the direction ofsearching flights of the ovipositing females. Obviously, this subject cannot be pursued at present beyond these qualitative remarks. Data for a quantitative evaluation of these influ­ences are lacking. In closing this discussion, however, it is worth pointing ou t that, if further work should confirm the hypothesis that larval sur­vival is optimal on shoots of sizes preferred for oviposition (p. 539), dif­ferential survival could be said to enhance the influence of habitat selec­tion on the distribu tion of Lipara lucens. X. SUMMARY I. This study concerns the analysis of some aspects of habitat selection by the chloropid fly Lipara lucens Mg., based on field data and the results of experiments done in the field and in the laboratory. II. The flies emerge in late spring and lay eggs on fresh shoots of Com­mon Reed (Phragmites communis Trin.). Immediately after hatching, the larvae migrate on the surface of the reed shoot to the top and then down­wards through the centre of the roll of young leaves until they reach a position just above the growing point. In the following period, during which the larva remains in the same place and feeds on the very young leaves, a gall is formed because the internodes of the stem remain short and grow in width. After completion of the gall in late summer, the larva eats its way through the growing point to reach the parenchyma­tous pith of the gall, in which it eats out a chamber. The larva hibernates in this gall chamber. III. The habitat ofLipara lucens has been characterized in the literature as reed vegetation on dry soil. DOCTERS VAN LEEUWEN assumed that the water level influences the gall distribution not directly but via the development of the reed shoots, which is less vigorous in dry than in wet conditions. This hypothesis was taken as a starting point for the present investigation. The criterion taken for shoot development was the basal diameter of the reed shoots, since this is the only easily measured pro­perty that remains reasonably constant throughout the season. IV. A partial regression analysis of the series of samples from three reed vegetations revealed that of three environmental variables, i.e. mean basal shoot diameter, shoot density, and water level, the partial regres­sion of shoot diameter on gall density was significant in all three cases. l I 546 J. H. MOOK No significant partial regression was found for shoot density, and partial regression ofwater level on gall density was significant in one series only. The correlation between shoot diameter and gall density was found to be caused both by a preference of the flies when laying eggs and by differential mortality of the larvae before gall completion. V, VI. Egg-laying was investigated in choice experiments done in a field cage on whole shoots and in the laboratory on top parts of shoots. In both situations a preference was found for shoots in the medium range of shoot diameters. Detailed observation of the behaviour of the flies in the laboratory showed that the numbers ofvisits to shoots ofall diameters were similar but that behaviour on the shoots differed, leading to longer visits and the laying of more eggs on shoots of medium diameter. Other laboratory experiments showed that shoot diameter plays a role as a stimulus, but that other stimuli, possibly chemical, are also im­portant. When the flies had access to shoots of only one diameter class at a time, the egg production on shoots of medium diameter remained greater than that on thicker shoots. VII. Survival of the larvae up to gall formation ""as studied in a field experiment. Survival was greatest on the thinnest shoots and decreased wirh increasing shoot diameter. No correlation was found between sur­vival and the distance the larvae had to travel on the surface of the shoot (which tends to be greater for the thicker shoots), despite the fact that the larvae are very vulnerable to drought. This phenomenon is probably explained by the fact that the larvae hatch during the night and are able to cover the distance wi thin a few hours, at a time when the shoot surface is moist with dew. Differential survival could be demonstrated for the next period, dur­ing which the larvae migrate from the surface near the top of the shoot to the growing point. Fewer larvae reach the growing point in thick shoots than in thin ones, probably because thick shoots have more layers of leaf sheaths to be passed and the distance from the top to the growing point is greater. Nothing is known about the interaction between larva and reed plant that results in gall formation. Combination of the estimates of egg distribution and survival in rela­tion to shoot diameter gives a probability distribution of galls on shoots of different diameters which resembles the average distribution of galls in the field rather well. It is remarkabl e that the prefcrence of the flies and the survival up to gall formation do not fall in the same classes, the latter being greater on shoots that are thinner than those on which most eggs are laid. HABITAT SELE VIII. Survival of the larva meter. Survival increases w effect of predation (especii and parasitization by thfj greater toll on the thinneJ braconid Polemon liparae, ill thicker shoots than on the most populations its ave rag terbalance the influence of IX. The distribution of egg period over shoots of differ the extent to which egg-layl conclusions can be drawn b in the estimate ofsurvival, 1 ference and survival clo nc additional factors are disc1 correspondence. The results of the anal. suggest an explanation of under natural conditions. ALLAN,S.E. and W. H. PEARSA! Phragmites. Oikos 14, 176- 1 AUBERT,]. F. and N. SHAUMAR, sexe chez les Ichneumonid, des cocons pleins c t des coco 2194--2195. AVJDov,Z ., lVLJ.Bl!:RLlNGER an< host specificity in the Brucl Callosobruchus chinensis L., Al BAKKER, D ., 1957; De levcnswijz riet in de l'<oordoostpolder. BAKKER, D., 1960; H e t botanisch 63- 79. BLAIR,K. G., 1932; Some notes dae). Entomologist's mono J BLAIR,K. G., 1944; Further note pidae). Entomologist's mon CAIN, A.]. and P. M. SHEPPARD, ton 39, 89-116. CLAUSEN, C.P., 1939; The effect parasites a nd its relation t( Soc. 47, 1-9. x 547 HABITAT SELECTION BY LIPARA L CENS MG. VIII. Survival of the larvae in the gall is also correlated with shoot dia­meter. Survival increases with increasing shoot diameter, as a combined effect of predation (especially by birds, i.e. Blue Tits, Parus caemleus) and parasitization by the chalcid Stenomalina lipame, which take a greater toll on the thinner shoots. The other important parasite, the braconid Polemon liparae, induces a somewhat higher mortality on the thicker shoots than on the thinner ones when it is present alone, but in most populations its average influence is not important enough to coun­terbalance the influence of the other two factors mentioned. IX. The distribution of egg laying and survival during the whole larval period over shoots of different diameters were compared to determine the extent to which egg-laying preference has survival value. No reliable conclusions can be drawn because of the accumulation of random errors in the estimate of survival, but the calculated curves for oviposition pre­ference and survival do not seem to coincide closely. However, some additional factors are discussed, inclusion of which may improve the correspondence. The results of the analysis of oviposition preference and survival suggest an explanation of the observed distribution of Lipara lucens under natural conditions. XI. REFERENCES ALLAN, S. E. and W. H . PEARSALL, 1963: Leaf analysis and shoot production in Phragmites. Oikos 14, 176-189. AUBERT,J. F. and N.SHAUMAR, 1962 : Nouvelle experience sur Ie determinisme du sexe chez les Ichneumonidae Pimplinae (Insectes hymenopteres): ponte dans des cocons pleins et des cocons creux. C.r. hebd . Seanc. Acad. Sci., Paris 225, 2194-2195. AVIDOV, Z., M.J. BERLINGER and S. W. ApPLEBAUM, 1965 : Physiological aspccts of; host specificity in the llruehidac. III Effect of curvature and surface al'ca on Callosobruchus chinensis L., Anim. Behav. 13, 178- 180. BAKKER, D., 1957: De Icvenswijzc van her rie t, in: D. Bakker en D. T. Bieuwinga: Het riet in de Noordoostpolder. Van Zec tot Land 21, 5-25. BAKKER,D., 1960: Het botanisch onderzock in de IJsseimccrpolders. Vakbl. BioI. 40, 63- 79. BLAIR, K. G ., 1932: Some notes on the galls of L i/Jarrt lua/ls :VIg. (Diptera, Chloropi­dae). Entomologist's mon oMag. 68, 10-· 13. BLAIR, K. G., 1944 : Further notes on the galls of Lij!ara hums Nfg. (Diptera, Chloro­pidae). Entomologist's mono Mag. 80,6-7 . CAIN,A.J. and P. M . SHEPPARD, 1954: Natural selection in Cepaea. Genetics, Prince­ton 39, 89- 116. CLAUSEN, C. P., 1939: The effect of host size- upon the- sex ratio of h ymenopterous parasites and its relation to methods of rearing and colonization. JI. N.Y. enl. Soe. 47, 1-9. 

Habitat selection by <i>Lipara lucens</i> Mg. (Diptera, Chloropidae) and its survival value

University of Groningen Research Database

  
 University of Groningen
Habitat selection by Lipara lucens Mg. (Diptera, Chloropidae) and its survival value
Mook, Jacob Hendrik
IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from
it. Please check the document version below.
Document Version
Publisher's PDF, also known as Version of record
Publication date:
1967
Link to publication in University of Groningen/UMCG research database
Citation for published version (APA):
Mook, J. H. (1967). Habitat selection by Lipara lucens Mg. (Diptera, Chloropidae) and its survival value
Groningen: Noordhoff Uitgevers
Copyright
Other than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of the
author(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons).
Take-down policy
If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately
and investigate your claim.
Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons the
number of authors shown on this cover page is limited to 10 maximum.
Download date: 10-02-2018
545 HABITAT SELECTION BY LIPARA LUCENS MG. 
This tendency to cluster, however, will be counteracted to some extent 
by the randomness in the direction ofsearching flights of the ovipositing 
females. 
Obviously, this subject cannot be pursued at present beyond these 
qualitative remarks. Data for a quantitative evaluation of these influ­
ences are lacking. In closing this discussion, however, it is worth pointing 
ou t that, if further work should confirm the hypothesis that larval sur­
vival is optimal on shoots of sizes preferred for oviposition (p. 539), dif­
ferential survival could be said to enhance the influence of habitat selec­
tion on the distribu tion of Lipara lucens. 
X. SUMMARY 
I. This study concerns the analysis of some aspects of habitat selection 
by the chloropid fly Lipara lucens Mg., based on field data and the results 
of experiments done in the field and in the laboratory. 
II. The flies emerge in late spring and lay eggs on fresh shoots of Com­
mon Reed (Phragmites communis Trin.). Immediately after hatching, the 
larvae migrate on the surface of the reed shoot to the top and then down­
wards through the centre of the roll of young leaves until they reach a 
position just above the growing point. In the following period, during 
which the larva remains in the same place and feeds on the very young 
leaves, a gall is formed because the internodes of the stem remain short 
and grow in width. After completion of the gall in late summer, the 
larva eats its way through the growing point to reach the parenchyma­
tous pith of the gall, in which it eats out a chamber. The larva hibernates 
in this gall chamber. 
III. The habitat ofLipara lucens has been characterized in the literature 
as reed vegetation on dry soil. DOCTERS VAN LEEUWEN assumed that the 
water level influences the gall distribution not directly but via the 
development of the reed shoots, which is less vigorous in dry than in wet 
conditions. This hypothesis was taken as a starting point for the present 
investigation. The criterion taken for shoot development was the basal 
diameter of the reed shoots, since this is the only easily measured pro­
perty that remains reasonably constant throughout the season. 
IV. A partial regression analysis of the series of samples from three reed 
vegetations revealed that of three environmental variables, i.e. mean 
basal shoot diameter, shoot density, and water level, the partial regres­
sion of shoot diameter on gall density was significant in all three cases. 
l 
I 
546 J. H. MOOK 
No significant partial regression was found for shoot density, and partial 
regression ofwater level on gall density was significant in one series only. 
The correlation between shoot diameter and gall density was found 
to be caused both by a preference of the flies when laying eggs and by 
differential mortality of the larvae before gall completion. 
V, VI. Egg-laying was investigated in choice experiments done in a field 
cage on whole shoots and in the laboratory on top parts of shoots. In 
both situations a preference was found for shoots in the medium range 
of shoot diameters. Detailed observation of the behaviour of the flies in 
the laboratory showed that the numbers ofvisits to shoots ofall diameters 
were similar but that behaviour on the shoots differed, leading to longer 
visits and the laying of more eggs on shoots of medium diameter. 
Other laboratory experiments showed that shoot diameter plays a 
role as a stimulus, but that other stimuli, possibly chemical, are also im­
portant. When the flies had access to shoots of only one diameter class 
at a time, the egg production on shoots of medium diameter remained 
greater than that on thicker shoots. 
VII. Survival of the larvae up to gall formation ""as studied in a field 
experiment. Survival was greatest on the thinnest shoots and decreased 
wirh increasing shoot diameter. No correlation was found between sur­
vival and the distance the larvae had to travel on the surface of the shoot 
(which tends to be greater for the thicker shoots), despite the fact that 
the larvae are very vulnerable to drought. This phenomenon is probably 
explained by the fact that the larvae hatch during the night and are able 
to cover the distance wi thin a few hours, at a time when the shoot surface 
is moist with dew. 
Differential survival could be demonstrated for the next period, dur­
ing which the larvae migrate from the surface near the top of the shoot 
to the growing point. Fewer larvae reach the growing point in thick 
shoots than in thin ones, probably because thick shoots have more layers 
of leaf sheaths to be passed and the distance from the top to the growing 
point is greater. 
Nothing is known about the interaction between larva and reed plant 
that results in gall formation. 
Combination of the estimates of egg distribution and survival in rela­
tion to shoot diameter gives a probability distribution of galls on shoots 
of different diameters which resembles the average distribution of galls 
in the field rather well. It is remarkabl e that the prefcrence of the flies 
and the survival up to gall formation do not fall in the same classes, the 
latter being greater on shoots that are thinner than those on which most 
eggs are laid. 
HABITAT SELE 
VIII. Survival of the larva 
meter. Survival increases w 
effect of predation (especii 
and parasitization by thfj 
greater toll on the thinneJ 
braconid Polemon liparae, ill 
thicker shoots than on the 
most populations its ave rag 
terbalance the influence of 
IX. The distribution of egg 
period over shoots of differ 
the extent to which egg-layl 
conclusions can be drawn b 
in the estimate ofsurvival, 1 
ference and survival clo nc 
additional factors are disc1 
correspondence. 
The results of the anal. 
suggest an explanation of 
under natural conditions. 
ALLAN,S.E. and W. H. PEARSA! 
Phragmites. Oikos 14, 176- 1 
AUBERT,]. F. and N. SHAUMAR, 
sexe chez les Ichneumonid, 
des cocons pleins c t des coco 
2194--2195. 
AVJDov,Z ., lVLJ.Bl!:RLlNGER an< 
host specificity in the Brucl 
Callosobruchus chinensis L., Al 
BAKKER, D ., 1957; De levcnswijz 
riet in de l'<oordoostpolder. 
BAKKER, D., 1960; H e t botanisch 
63- 79. 
BLAIR,K. G., 1932; Some notes 
dae). Entomologist's mono J 
BLAIR,K. G., 1944; Further note 
pidae). Entomologist's mon 
CAIN, A.]. and P. M. SHEPPARD, 
ton 39, 89-116. 
CLAUSEN, C.P., 1939; The effect 
parasites a nd its relation t( 
Soc. 47, 1-9. 
x 
547 HABITAT SELECTION BY LIPARA L CENS MG. 
VIII. Survival of the larvae in the gall is also correlated with shoot dia­
meter. Survival increases with increasing shoot diameter, as a combined 
effect of predation (especially by birds, i.e. Blue Tits, Parus caemleus) 
and parasitization by the chalcid Stenomalina lipame, which take a 
greater toll on the thinner shoots. The other important parasite, the 
braconid Polemon liparae, induces a somewhat higher mortality on the 
thicker shoots than on the thinner ones when it is present alone, but in 
most populations its average influence is not important enough to coun­
terbalance the influence of the other two factors mentioned. 
IX. The distribution of egg laying and survival during the whole larval 
period over shoots of different diameters were compared to determine 
the extent to which egg-laying preference has survival value. No reliable 
conclusions can be drawn because of the accumulation of random errors 
in the estimate of survival, but the calculated curves for oviposition pre­
ference and survival do not seem to coincide closely. However, some 
additional factors are discussed, inclusion of which may improve the 
correspondence. 
The results of the analysis of oviposition preference and survival 
suggest an explanation of the observed distribution of Lipara lucens 
under natural conditions. 
XI. REFERENCES 
ALLAN, S. E. and W. H . PEARSALL, 1963: Leaf analysis and shoot production in 
Phragmites. Oikos 14, 176-189. 
AUBERT,J. F. and N.SHAUMAR, 1962 : Nouvelle experience sur Ie determinisme du 
sexe chez les Ichneumonidae Pimplinae (Insectes hymenopteres): ponte dans 
des cocons pleins et des cocons creux. C.r. hebd . Seanc. Acad. Sci., Paris 225, 
2194-2195. 
AVIDOV, Z., M.J. BERLINGER and S. W. ApPLEBAUM, 1965 : Physiological aspccts of; 
host specificity in the llruehidac. III Effect of curvature and surface al'ca on 
Callosobruchus chinensis L., Anim. Behav. 13, 178- 180. 
BAKKER, D., 1957: De Icvenswijzc van her rie t, in: D. Bakker en D. T. Bieuwinga: Het 
riet in de Noordoostpolder. Van Zec tot Land 21, 5-25. 
BAKKER,D., 1960: Het botanisch onderzock in de IJsseimccrpolders. Vakbl. BioI. 40, 
63- 79. 
BLAIR, K. G ., 1932: Some notes on the galls of L i/Jarrt lua/ls :VIg. (Diptera, Chloropi­
dae). Entomologist's mon oMag. 68, 10-· 13. 
BLAIR, K. G., 1944 : Further notes on the galls of Lij!ara hums Nfg. (Diptera, Chloro­
pidae). Entomologist's mono Mag. 80,6-7 . 
CAIN,A.J. and P. M . SHEPPARD, 1954: Natural selection in Cepaea. Genetics, Prince­
ton 39, 89- 116. 
CLAUSEN, C. P., 1939: The effect of host size- upon the- sex ratio of h ymenopterous 
parasites and its relation to methods of rearing and colonization. JI. N.Y. enl. 
Soe. 47, 1-9. 


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Habitat selection by <i>Lipara lucens</i> Mg. (Diptera, Chloropidae) and its survival value

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