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Page 1: A SNAIL-COLLECTING APHIS-LION LARVAw3.marietta.edu/~biol/378/snail_collecting_aphis_lion...A SNAIL-COLLECTING APHIS-LION LARVA By DAVID T. JONES Research Laboratory of Biology, Marietta
Page 2: A SNAIL-COLLECTING APHIS-LION LARVAw3.marietta.edu/~biol/378/snail_collecting_aphis_lion...A SNAIL-COLLECTING APHIS-LION LARVA By DAVID T. JONES Research Laboratory of Biology, Marietta

A SNAIL-COLLECTING APHIS-LION LARVABy DAVID T. JONES

Research Laboratory of Biology, Marietta College

On October 27th, 1928, while on the Hocking River Expedition(a cooperative expedition sent out by Marietta College and the OhioBiological Survey to further the state molluscan survey), the authordiscovered an insect larva bearing on its back a number of minutesnail shells. Professor H. R. Eggleston and Mr. Paul Crone werecalled to observe the form before it was killed. Diligent search ofthe immediate vicinity resulted in a collection of five specimens, allof which were preserved in 80 per cent. alcohol.

The find was made while searching for minute snails under moistleaves on a hillside covered chiefly with hardwood trees. The areain which the specimens were found could not have been more thansix feet in diameter. The larvae were active enough to cause alert-ness on the part of the collector. The weather was cloudy with fre-quent showers. The location was roughly estimated as seven mileswest of Albany, Ohio, and twelve miles east of MeArthur, in theeastern part of Vinton County. The author had, a few weeksbefore, observed a similar larva bearing snails in Squaw Hollownear Marietta, but the specimen escaped capture.

In classification the larva seems to belong either to the familyCh1-ysopidae or the family H emerobiulae, probably in the former.As yet the research monographs relating to the group have not beencollected. Restriction of the latter family in recent years has madeolder classifications worthless. Many classifications depend onadult characteristics instead of larval characteristics, necessitatinga knowledge of more facts of the life history than we have at pres-ent collected. There seems to be some discrepancy between Englishand American authors as to larval characteristics. According toImms (1925), this larva certainly goes in the family Chrysopidae,because it has the setae arising from dorso-lateral tubercles. Thepattern on the dorsal part of the head and the differentiation of thedistal part of the antennae compare favorably with the figure (fig.316, p. 470) of Ch1'ysopa paZZida as given by Sharp after Brauerin the Cambridge Natural History. Habitat and known occurrenceof adult forms point to itsbeing rather a chrysopid than a hemero-biid. It is hoped that observations in the field and literature now

1

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2 A SNAIL-COLLECTING APHIS-LION LARVA

being collected here will add to the known facts of its life historyand secure its definite classification. The aim of the present articleis to study the external morphological features.'

The most noticeable feature about the aphis-lion larva is the bur-den or cloak on its back. Aphis-lion larvae usually feed on aphidsor plant lice. After destroying the plant lice by thrusting theirsucking spears into them and sucking out the life-juices, the aphis-lions place what is left of the bodies of their victims on their backs,where they are held by hooked hairs and tufts of setae or bristles.Other debris is usually mixed with the aphis hulls. This species,however, differs in that the burden was chiefly snail shells and insecthulls. One specimen had the following on its back:

Insect remains:I tibia, probably of Orthopiero:I head of a small insect, perhaps a cricket.I head of another small insect.2 abdomen-hulls of small beetles of different species.2 separate, spiny, thoracic segments of some insects, both

segments probably from the same species.I thoracic hull of a larger insect.5 unidentifiable fragments of insect skeletons.

Snail remains:5 Punctum pygmaeum (Drap.j-i-I adult and 4 juvenile.

The adult had a portion of the liver remaining in theshell. Two juveniles had the liver and apparently thebody intact within the shell and may have been livingwhen preserved. All other shells of this species wereempty. One was broken, apparently an old shell whenit was picked up.

I Euconulus fulvus (Muller r=-A four-whorled juvenile. Asmall hole in the upper part of the third whorl from theapex was about the size and shape.of the cross-sectionof the sucking spears of the larva. As this shell is thin,it is possible, though not probable, that the snail wasthus killed. The shell was empty.

I juvenile, unidentifiable, nuclear whorls alone present.Not a large form.

I Striatura milium (lVIorse)-Empty shell though notweathered. I have not taken this previously in myOhio collections.

The largest shell, the juvenile Euconulus, was under 2 mm. 111

diameter.

Inasmuch470) noticedand placed irbles and beninasmuch asclothes itselfare very difffeasibility ofrobbing therrlecting at thione juvenileit took the la

The shell!web-like struof the shell (the case of !some twistintough and elOnly two sfive specimer

No oneAnthony (Cneuropteranthe fresh watary tract issecreted by tThis silk is tdIe intestinewhence it i:dilated rectaneret. Thein many habthe aphis-lioas Dewitz 01the threadsthreads conithis as imprthe shell neshooked hain

The burlof the midlispecies the

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- DAVID T. JONES J

life historyesentarticle

Inasmuch as "Dewitz (Sharp, Cambridge Natural History, p.470) noticed that some specimens that he denuded of their clothingand placed in a glass, seized small bits of paper with their mandi-bles and bending the head, placed morsels, on their backs"; andinasmuch as this species instead of wearing the usual aphis hulls,clothes itself chiefly with the smallest snails in this vicinity whichare very -difficult to obtain, the snail collector wonders as to thefeasibility of domesticating some of these larvae and persistentlyrobbing them while on collecting trips. In fifteen minutes of col-lecting at this locality, before finding the larvae; I had 'found onlyone juvenile specimen of Punctum pygmawm. I wonder how longit took the larva to find the five!

The shells and hulls are held in place by twisted strands of cob-web-like structure. These mats are applied usually to only one sideof the' shell or object; frequently on the side of the body whorl inthe case of snails. The strands run irregularly, though there issome twisting to form stouter strands. These strands are verytough and elastic, even after remaining several months in alcohol.Only two shells and 'a few insect fragments came off of thefive specimens while they were being brought back to the college.

No one knows how this spider-like silk is secreted. MissAnthony (Comstock, pp. 282-283) in her investigations of theneuropteran genus Sisyra, the spongilla flies, whose larvae feed onthe fresh water sponge, finds that the posterior part of the alimen-tary tract is modified into a silk-producing mechanism, silk beingsecreted- by the modified middle portions of the Malpighian tubules.This silk is thrown into the posterior portion of the atrophied mid-dle intestine, which is enlarged to form a silk-receptacle, -fromwhence it is conveyed through a narrowed hind intestine to adilated rectal reservoir, from whence it is spun from- an anal spin-neret. The aphis-lion differs from the spongilla fly in habitat andin many habits, even though the two are in neighboring genera. Ifthe aphis-lion larva places objects on its back with its mouth partsas Dewitz observed, it would be interesting to observe how it spinsthe threads to hold them. Sharp in discussing similar hemerobiidthreads considers that they might be spiders' webs, but I regardthis as improbable because the attachment is .usually at the side ofthe shell nearest the larva. The web and not the shells cling to thehooked hairs of the back (q, v. in the following discussion).

The burden does not rest directly on the back. In the vicinityof the midline it is held up by hooked hairs, while laterally in thisspecies the tubercles are produced into pedicels which bear long

a is the bur-d on aphidsusting their;, the aphis-their backs,or bristles.

'his species,s and insect

eies,isects, both

4 juvenile.[ling in thearently thebeen livingpeeies wereshell when

ivenile. Arl from theross-sectionhell is thin,l snail was

te present.

nough notsly in my

r 2 mm. in

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4 A SNAIL-COLLEC'fING APHIS-LION LARVA

ment is wtapering iusually weannulationsfurcation itions. Thetenna beforof two smation consistleast twent

The eyenotch in throunded ee

shaped figApparentlythe centrallies undereye, and thother hand,of the presof the "gu

Ventral!ing spears.and a closeface of thesucking gring maxillaof this uepoint. Botwith the pIaction. Th!absorbed.the alimentmodified inmodifieatioexternal inThe labiuis a palpigspears. Tuniformlyfollow twesmall and i

setae, usually curving outward and upward, like the prongs of abundle-carrier on a binder, to hold up the burden. Apparently aircan circulate between the burden and the back.

The usefulness of the bundle is questionable. A shade-inhabit-ing larva does not need it for protection from the sun. The burdenis very conspicuous when the larva moves. It might serve to con-ceal the larva before it starts to run. It then appears to be a heapof empty shells, the" bleaching bones" of the" last stand" of somesnail colony whose retreat tragically terminated in some slowly dry-ing pocket, where the moisture eventually all evaporated. Such aheap of dead shells would appear very unattractive to anything oranybody, except the conchologist, and even he, startled at seeingsuch a pile of supposedly dead shells walking bodily away, mightallow the larva to escape. Considering all factors it is probablethat the burden is more a hindrance than a help. It may be but avestigial style of clothing or habit of adornment, that has persistedlong after it ceased to be useful. If this be true, the supportingpedicels are a most remarkable morphological adaptation to thesenses of beauty and propriety (whatever those may be in aphis-lion larvae). These long appendages, with their accompanyinghairs, hold this headgear (or rather backgear) in place and up offthe skin. This is more than any hatpin yet invented can do. Adescription of this apparatus will be given further on in this article.

Since the aphis-eating, aphis-lion larvae carry the graveyard oftheir aphis victims on their backs, further investigation should bemade to determine as to whether this species, in varying from theusual adornment of aphis hulls, has not varied from the usual dietof aphids.

The head of this larva is flattened dorso-ventrally and producedlaterally to form very broad eye peduncles. Ventrally it is a variedbuff like the remainder of the animal, but dorsally the head exhibitsthe following color markings: a buff anchor in mid-line, the handleof which forks like a Y, and is directed anteriorly. Lateral to eachprong of the Yare two intense black dashes, extending transversely.A triangular pigmented area lies within the forks of the Y. Therest of the dorsal part of the head is lightly pigmented, except themargin which is lighter. Laterally a lightly pigmented patchoccurs some distance posterior to the eye. A labrum projects ante-riorly over the labium. Dorsally the head bears a pair of antennaeof a filiform type, each of which on close observation is seen to bebranched. Each antenna sits in a socket which rises above the sur-face of the head dorsal to the ends of the labrum. The first seg-

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DAVID T. JONES 5

prongs of aparently air

ment is well-defined, large and stout; the second, elongate, buttapering into those more distad. Then follow fourteen short,usually well-defined segments, which grade abruptly into mereannulations of which there are fifteen, more or less, until the firstfurcation is reached. This branch consists of three small annula-tions. There are approximately thirty-six annulations on the an-tenna before the second furcation is reached. This branch consistsof two small annulations. The antenna distal to the second furca-tion consists of a very fine tapering thread, estimated to contain atleast twenty-five 'annulations,

The eyes are laterally located. They are elliptical except for anotch in the ventral side of each. Each consists of five peripheralrounded "openings" grouped around a central sixth. The star-shaped figure thus formed resembles the guard of a wrist watch.Apparently some circular dark body, which can be seen throughthe central "hole" and partially through the peripheral "holes,"lies under this "guard" in the center. Perhaps this is the real~eye, and the pigmented "guard" is a protective device. On theother hand, the openings may be sunken facets, shrunken by actionof the preservative, and the" underlying eye" may be but shadowsof the "guard. ' ,

Ventrally the head is smooth except for the supports of the suck-ing spears. Each sucking spear is composed of a dorsal mandibleand a close-fitting maxilla that fits in a groove on the ventral sur-face of the former. The latter has a basal stipes and cardo, Asucking groove is formed between each mandible and its correspond-ing maxilla. The end of each sucking spear consists of the openingof this sucking tube, and a mesial inner projection, the piercingpoint. Both sucking grooves at their proximal ends communicatewith the pharynx, which in this group is said to have a pumpingaction. The mouth is said to be vestigial and the liquid food allabsorbed. The vestigial mouth and the vestigial posterior part ofthe alimentary canal, into which wastes never come and which ismodified in Sisyra into a silk-producing apparatus, are evidentlymodifications correlated with the sucking of liquid foods. Allexternal indications point toward similar modifications in this form.The labium is smooth and at its corners it bears what apparentlyis a palpiger. The labial palps are almost as long as the suckingspears. They are clavate in shape and hairy. Segmentation is notuniformly distinct. The basal segment is large and distinct. Thenfollow twenty-one (more or less) segments which are proximallysmall and indistinct. These end distally with a wide-spreading seg-

ade-inhabit-The burdenerve to con-to be a heapid" of someslowlydry-

ed. Such aanything or~dat seeingrway, mightis probable

lay be but aias persistedsupporting

rtion to thebe in aphis-companying~and up offcan do. Athis article.raveyard ofn should be19 from thee usual diet

ld producedt is a variedead exhibits, the handleeral to eachransversely.he Y. The, except the:nted patch-ojects ante-of antennae3 seen to beove the sur-he first seg-

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6 A SNAIL-COLLECTING ArHIS-LlON LARVA

ment, into which the last portion of the palp is inserted. This lastportion at its base consists of five very minute annulations, but itsdistal part is smooth.

The prothorax is triply divided dorsally. The mesothorax andmetathorax consist of but one lobe, each dorsally. Each thoracicsegment bears a pair of elongate pedicels which in turn bear setae.These support the burden. The pedicels are inserted in the bodyjust dorsad to the leg sockets. Each thoracic pedicel is flattenedand slightly enlarged at its distal end. The distal end bears fouror five pairs of hairs, the members of a pair being situated dorso-ventrally. The ventral hairs are short and needle-like, but thedorsal hairs are long and coil in tendril fashion at their distal ends.Along each side of a pedicel are usually eight or nine single hairs,usually of the long, coiling type; frequently interlocked with sim-ilar hairs on neighboring pedicels. Hairs may occur on the dorsalsurface of a thoracic pedicel, but the ventral surface is usually bare.The prothoracic pedicels, situated on the middle lobe of the pro-thorax, are directed forward. Both mesothoracic and metathoracicpedicels are directed laterally.

Of the ten abdominal segments, six (segments two to seven in-clusive) bear fully developed, paired, abdominal pedicels. Aspiracle occurs on each one of these segments just dorsal to the baseof each pedicel. Segment eight has the rudiments of a pedicel anda fully developed spiracle. Abdominal segment one has a spiracle,but no pedicel. The posterior prothoracic lobe bears a prothoracicspiracle on each side. Abdominal pedicels differ from thoracic inthat most of the setae are borne at the tips of the pedicel. Thedorsal side of the pedicels may bear smaller hairs. The setae onthe abdominal pedicels are usually three to four of the long-hookedtype and three to four of the short needle-like kind, the latteroccurring ventral to the. former. 'I'he abdominal appendages slantprogressively caudad the farther posteriorly they are situated.

The hairs of the back are the same as those on the pedicels, ex-cept finer. They are hooked or with coiled ends, the coiling usuallydirected posteriorly. Close examination of the dried end-coils ofthese latter leads one to advance the theory that the "silk" whichbinds the shells together in this animal may be only entangled pro-longations of certain of these hairs. The hairs of the back arearranged in transverse rows, as follows:

(1) On the prothorax=-Two prominent stout bristles betweenthe bases of the prothoracic pedieels.

(2) Onh

(3) On1'1

(4) Onmaba

seSegment

and in Sisytigial in thspigots ofjointed. T

On thesockets therparatively Itive thoracirelatively shairs, and ecurved tarsseeminglyawith the staorgan of usuction. Tproximal er

AXTHONY, M!Vol.

BANKS, NATI

Soc.,Bl;AUER, FRED

COMSTOCK, J1925

CONSTANT an1881

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DAVID T. JONES 7

a. This lastI. b .nons, ut Its

(2) On the mesothorax-Two stout bristles and four smallerhairs between bases of pedicels.

(3) On the metathorax-Ten or more hairs on a transverseridge between bases of pedicels.

(4) On abdominal segments-In the first seven abdominal seg-ments, two rows of from twelve to fifteen hairs extendacross each segment; one of these rows is on the ridgebetween the bases of the pedicels (in first segment thereare no pedicels but hairs are there), the other row occursin the groove between segment ridges. These rows be-come less defined posteriorly. The last three segmentshave fewer hairs, though there are still indications ofrows. On segment nine, which is elongate, a row oftransverse hairs OJl a ridge make it look much like twosegments.

Segment ten of the abdomen bears the anus which is vestigial,and in Sisyra, transformed into a spinneret. It appears to be ves-tigial in this larva also. Two ventral anal projections resemblespigots of a spider's spinneret in shape, though they are notjointed. Two lateral expansions of the anus also occur.

On the outer side of both mesothoracic and metathoracic legsockets there is a pore of some kind. The coxae of the legs are com-paratively large and are set in sockets projecting from their respec-tive thoracic segments. The trochanters are distinct; the femursrelatively small; the tibias slender, each with four rows of spinyhairs, and each tarsus has a single joint. Each tarsus ends in twocurved tarsal claws, between which is a sac-like empodial base withseemingly a glandular surface. A stalk of this sac connects distallywith the stalk of the empodium proper, which is a trumpet-shapedorgan of unknown function, apparently built on the principle ofsuction. Two short, annulated hairs are attached beneath theproximal end of the empodial base.

othorax andach thoracic

bear setae.in the bodyis flattened

d bears fouruated dorso-ike, but ther distal ends,single hairs,

ed with sim-on the dorsalusually bare.

of the pro-metathoracic

to seven in-pedicels. AIto the basepedicel and

as a spiracle,a prothoracic

thoracic inedicel. The

The setae onlong-hooked

d, the latterndages slant

·ituated.pedicels, ex-iling usuallyend-coils of

"silk" whichtangled pro-

the back are

BIBLIOGRAPHY

lstles between

AX'l'llOXY, MAUD H.-The Metamorphosis of Sisy1'O, The American Naturalist,Vol. 36, 1902 (unavailable).

BAXKS, NATHAN-A Revision of the Nearctie Hemerobiidae, Trans. Am. Ent.Soc., Vol. 32, pp. 21-51, 1905 (unavailable).

BIlAUER, FREDERlcH-Die Neuropteren Europas, Wien, 1876 (unavailable).COMSTOCK, JOHN HENRy-An Introduction to Entomology, Second Edition,

1925.CONSTANT and LUCAS-Bull. Soc. Ent. France (6), i, pp. XXI and XXXI,

1881 (unavailable),

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8 A S AIL-COLLECTING APHIS-LION LARVA

DEWITZ, H.-BioI. Centralblatt, IV, 'P. 722, 1885 (unavailable).I)IMS, A. D.-A General Textbook of Entomology, Methuen and Co., Ltd., 30

Essex St., W. C., London, 1925.SHARP, DAVID-Insects, Part I, The Cambridge Natural History, Vol. 5, Mae-

millan, 1895.S)UTH, R. C.-A Study of the Biology of the Ch1"ysopidae, Ann. Ent. Soc. Am.,

Vol. 14, pp. 27-35 (unavailable).WARD, HENRY B. and 'WHIPPLE, GEORGEC.-Fresh Water Biology, 'Wiley and

Sons, 1918.

Fw.12

I'

KEY TO PLATE I FIG. 7.

FIG. 1. Lateral view of aphis-lion larva.a. Burden or cloak.b. Right prothoracic pedicel.c. Seta or hair.d. Last left abdominal pedicel.e. Right antenna.f. Right sucking spear.g. Left labial palp.h. Last abdomiual spiracle.i. Eye.

KEY TO PLATE IIFIG. 2. Right metathoracic pedicel, ventral view.

a. The pedicel.b. Seta or hair.c. Seta from mesothoracic pedicel.d. Seta from abdominal pedicel.

FIG. 3. Ventral view of third left abdominal pedicel.a. Coiled seta or hair.b. Ventral needle-like hair.c. The pedicel.d. Seta of second left abdominal pedicel.

FIG. 4. Dorsal view of abdominal somites nine and ten.a. Segment nine.b. Vestigial anus.c. Anal projections.

FIG. 5. Dorsal view of left antenna.a. Socket.b. First annulation (as contrasted with segments proximally).c. First furcation.d. Secon d furca tion.

FIG. 6. Ventral view of head (antennae omitted).a. Sucking spear.b. Piercing point.c. Sucking tube.d. Maxilla (beneath in groove).

FIG.

FIG. 9.

FIG. 10

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DAVID T. JONES

Co.,Ltd., 30 1e. Mandible (above).f. Stipes of maxilla.g. Cardo of maxilla.h. Labium.i. Labial palpus.j. Labrum.k. Eye.1. Palpiger.

, Vol. 5, Mac·

lEnt.Soc.Am.,

:gy, Wiley and

KEY TO PLATE IIIFIG. 7. Dorsal view of head and prothorax.

a. Buff anchor.b. Transverse dashes.c. Eye.d. Base of sucking spear.e. Base of antenna.f. Labrum.g. Prothorax.h. Base of prothoracie pedicel.i. Pro thoracic spiracle.j. Base of mesothoracic pedicel.k. Bristle or seta.

Side view of hair of back (arrow points posteriorly).Right eye.

a. Ventral notch.b. "Opening."c. Central" opening."d. "Underlying eye."e. " Guard. "

FIG. 8.

FIG. 9.

FIG. 10 and 11. Empodia as seen when viewing larva laterally.a. Ordinary hair on the tarsus.b. Tarsal claw.c. Annulated hair.d. Empodium.e. Empodial base.

FIG. 12. Left prothoracic leg.a. Coxa.b. Trochanter.c. Femur.d. Tibia.e. Tarsus.f. Tarsal claw.g. Empodium.

~s proximally).

9

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