The Life History and Cocoon Spinning Behaviour of a South African Mantispid (Neuroprera : Mantispidae)

J. L. BISSETT and V. C. MORAN

Department of Zoology and Entomology, Rhodes University, Grahamstown

The life histories of South African mantispids have not been documented and reports on mantispid biology are generally scarce. As stated by Milliron (1940) it is evident that prior to 1934 most of the information on the biology of this family was based on the classical experiments of Brauer (1869) who worked on a European species Mantispa styriaca Poda. A more recent paper on manrispids by McKeown & Mincham (1948) is an account of the biology of the Australian sp~ciesMantispa uittata Guerin. The reports of Main (1931), Eltringham (1932), Kasron (1938) and Hungerford (1939) do not advance our knowledge of th5;e animals significantly. In the present investigation the life history and cocoon spinning behaviour of a local mantispid was studied. Difficulty was experienced, however, in identifying this mantispid and Tjeder (1966) has said : "Before a rrliable identificarion of mantispid specimens from Africa can be done it is necessary toexamine the types of all themany described species. The late Prof. Handschin of Basel has some few- years ago dealt with African species in some large pzpers but I regret to say that I have not found it possible to determine my specimens from these papers only. I think the family is a very difficult one to work with, the variability within the species apparently being great and the genitalia affording less clear distinguishing characters than in most Neuroptera. I do not know of anyone who for the present may be able to determine your material". As a result positive identification of this South African mantispid has not been achieved. However, the animals used in this study were always associated with Cape Chestnut trees, Calodendrurn capense (L.s.) Thunb., and adult specimens have been stored in the National Collection of Insects, Plant Protection Research Institute, Pretoria under the accession number AcP 5469. This insect will be referred to as the chestnut mantispid in this account. METHODS Methods used for rearing the larvae of the chestnut mantispid were simple but effective and similar to those used by McKeown & Mincham (1948) for rearing M. uittata. The first instar triungulin larvae were placed in a hollowed cork together with suffi- cient spiders' eggs or immature spiders as food. The larvae were observed every 24 hours and measurements taken across the widest point of the body. The animals were maintained at laboratory temperatures (March to June 1966, approximately 15'-20°C.) but at a high humidity which was achieved by enclosing the corks with the larvae in petri-dishes, the bottoms of which were lined with moist filter paper. In the chestnut Bissett and Moran : A South African mantispid 83 mantispid there are only three larval instars and the immature stages feed on the eggs of any spiders and may feed on immature spiders. Even larvae which had fed on the eggs of a single spider species for almost the whole larval life would feed readily when presented with the eggs of a different spider. This lack of host specificity in the chestnut mantispid is in agreement with the findings of several workers (Brauer, 1869, and Poujade, 1898, both cited by Milliron, 1940; Main, 1931; Smith, 1934; Kaston, 1938; and McKeown & Mincham, 1948). Some mantispids however are known to parasitize coleopterans (Werner & Butler, 1965); noctuid pupae (Woglum, 1935, cited by Milli- ron, 1940); and Hymenoptera (Smith, 1863, cited by Milliron, 1940).

LIFE HISTORY The life history of the chestnut mantispid may be briefly stated before dealing with each stage in more detail. The eggs take approximately 16 days to hatch and the first stage larvae on emergence are active, campodeiform triungulin larvae which feed on spiders' eggs. From 5-9 days after commencemenx of feeding the triungulin larva moults to a fatter more sluggish second stage larva and 2-5 days later the third stage is reached. By the end of the third stage hypermetamorphosis is complete and after 2-6 days of active feeding in this instar the bloated larva spins a cocoon and the prepupal stage is achieved. A period of 9-15 days quiescence follows before thepupa is formed and the adult emerges 20-28 days later. -" The eggs of the chestnut mantispid are deposited in circular batches on the uppcr surface of leaves of C. capense. Counts of 16 egg batches gave a maximal number of 1;650 eggs, a minimum of 2 14 and a mean of 836 eggs per batch. These figures compare with those of Smith (1934) who records a total of 2,200 eggs in six batches for Mantispa saoi Banks and with Clirnaciella brunnea var. accidentalis Banks in which a female deposited 1,028 eggs in a single batch (Hoffmann, 1936, cited by McKeown & Min- cham, 1948). Smith (1934) records that C. brunnea Say lays a total of only 250 eggs per female when in captivity. The eggs themselves (fig. la) are elongate, stalked. with a distinct micropylar cap at the anterior end and are a creamy yellow colour when first laid. The size of the eggs was fairly constant having a length of 0.38 mm and a width of 0.13 mm. Measure- ments of 57 egg stalks from different egg batches showed that they varied in length from 0.38 mm to 1.25 mm with a mean of 0.82 mm. The duration of the egg stage in the chestnut mantispid was only determined from one batch of eggs and was found to be 16 days. This agrees with the results of McKeown & Mincham (1948) from M. vittata where he found an egg duration of 16-18 days in warm weather.

First instar laroa Just prior to the emergence of the first instar larva the eggs become darker and the developing larvae can be seen clearly through the egg chorion (fig. Ib). At this stage the larva begins muscular "peristaltic" movements of the body and these movements cause the thoracic region of the larva to be pushed anteriorly thus exerting pressure at the point of rupture in the egg. The "peristaltic" movements cor~tinuefor three or four minutes at intervals of about eight seconds, until eventually the chorion of the egg ruptures. The movements persist and the dorsal part of the prothorax is pushed through the ruptured chorion followed by the head, which is reflexed beneath 84 3.ent. Soc. sth. Afr. Vol. 30, No. 1, 1967

Fig. 1. Chestnut mantispid egg showing the entire egg (la) and the larva in the egg just prior to emergence (lb). E-egg; La-larva; MC-micropylar cap; RPt-rupture point; St-stalk; StB-stalk base.

the thoracic segments. Once the chorion has ruptured, emergence is rapid taking between three and four minutes. The "peristaltic" movements force the last segments of the abdomen out of the egg. but the larva remains attached to the ruptured chorion by means of a caudal sucker. The larva now proceeds to free its head, which is attached to the sternite of the fourth abdominal segment by a transparent membrane. This is achieved by manipulating the head, mandibles and labial palpi and the head is freed in five or six minutes. The larva then remains motionless for some minutes, its caudal sucker still attached to the ruptured chorion of the egg and once the integument has hardened, the larva detaches itself. The newly emerged larva (fig. 2a-el, which measured approximately 1.1 mm in length, remains in close proximity to the other mantispid eggs for about two days before becoming active and starting to feed. In M. sbriaca (Brauer, 1869, cited by Milliron, 1940) and M. vittata (McKeown & Mincham, 1948) the newly emerged larvae hibernated for several months. This was not the case with the chestnut mantispid and this species died within 11 days if it had not found a suitable host. The method of location of a suitable host in the chestnut mantispid is obscure. The active triungulin larvae were not attracted to spiders eggs even if the host eggs were placed within a few centimetres of the larvae. The first stage Bissett and Moran : A South African mantispid

M x

MdGr

Fig. 2. Chestnut mantispid first instar triungulin larva showing some of the appendages. Pa-entire larva; 2b-left mandible and maxilla; 2c-right antenna; 2d-labial palpi; 2e-mesothoracic leg. Ant-antenna; CPr-cone like projection; CS-caudal sucker; F1-flagellum; Md-mandible; MdGr-mandibular groove; Mx-maxilla; Pdc-pedicel; Scp-scape; Tar-tarsus; TarC1-tarsal claw; Tb-tibia. 86 J. ent. Soc. sth. Afr. Vol. 30, KO.1, 1967 larvae are strong walkers and walking is characteristically accompanied by the head swaying from side to side and the caudal sucker being attached to the ground when the animal pauses. It is possible that host location is as a result of random wanderings on the part of the larvae. It is also a possibility that host location is aided by wind dispersal of the larvae or phoresy. Hungerford (1939) and Kaston (1938) report that mantispid triungulins may attach themselves to the bodies of female spiders and thus are possibly transported to the eggs or immature spiders. In any event once the first stage chestnut mantispid has located a host there is a marked increase in its activity. The larva moves around over the eggs stopping at intervals to apply the antennae, mandibles and labial palpi to the egg surface. About 24 to 48 hours later the pointed mandibles are closely applied to the surface of the spiders' egg and inserted by a series of upwardly directed movements of the head, each mandible then functioning as an individual sucking tube. The feeding triungulin larvae wrap their elongate bodies around the egg but the ventral surface of the bodv is kept away from the egg chorion by the long sternal setae which are present on abdominal segments two to seven. Observations on 24 individuals indicate that feeding continues for five to nine days before the triungulin moults to the second stage larva, during which time the abdomen of the triungulin takes on the colour of the spider's egg and becomes extremely bloated, increasing in width from 0.1 mm at the start of the instar to 0.5 mm at the end. The maximum widthrecorded was 0.9 mm and the minimum width 0.31 mm at the end of the instar.

Second and third instar larvae Approximately eight hours before moulting to the second instar the swollen triungulin larva ceases to feed, becomes inactive, and the cuticle becomes noticeably sticky. At the commencement of moulting the larval cuticle splits dorsally and longitudinally in the thoracic region. The head is reflexed beneath the ventral surface or'the thorax and the larva then performs "peristaltic" contractions which pass anteriorly up the body. These contractions enlarge the existing split and cause the exuviae to move slowly towards the posterior end of the body. As the exuviae moves posteriorly the head is withdrawn, followed by the thoracic and abdominal segments until eventually the exuviae falls away. The second stage larva feeds voraciously and showed a mean increase in width from 0.5 mm at the start of the instar to 0.9 mm at the end of the second stage; the minimum width measured at the end of the second instar was 0.75 mm and the maximum 1.1 mm out of a total of 16 measurements. The duration of the second instar was two to five days before moulting occured to the third stage larva (fig. 3a). The la~teris an extremely voracious feeder increasing in mean body width from 0.9 mm at the start of the instar to 2.1. mm at the end of the iristar. The minimum width measured at the end of the instar was 1.8 mm and the maximum 2.5 mm out of a total of 14 measurements. Hypermetamorphosis is complete at the end of the third stage, the legs are short and non-functional and the abdomen is extremely swollen. During. the third instar the action of the pharyngeal sucking pump can be seen clearly through the cuticle of the head and the animal can suck a spider's eqg dry within 20 minutes. Besides obvious differences in size between second and third instar larvae the latter are covered by prominent setae especially on the ninth and tenth abdominal segments and at the moult to the third stage larva the caudal sucker, which persists in the second stage, becomes modified as a spinneret. Spiracles are also clearly \,isible on the prothorax and first eight abdominal segments of the third stage larva. The duration of the th~rdinstar was from Bissett and Moran: A South African mantispid

, lrnrn ,

Fig. 3. Fully engorged third instar larva (3a) and pupa (3b) of the chestnut mantispid. An- anus; Ant-antenna; CpdE-compound eye; DevL-developing legs; Hks-hooks; Md-mandible; RptL-raptorial foreleg; Sp-spiracle; Spn-spinneret; WBd-wing bud. 88 J. ent. Soc. sth. Afr. Vol. 30, No. 1, 1967 two to six days and at the end of this instar the engorged larva constructed a silken cocoon in which the prepupa was formed. Larvae commenced cocoon spinning when the supply of eggs was exhausted irrespective of whether they were fi.111~engorged or not.

Prepupal and pupal stage After the cocoon has been completed the larva enters a prepupal period which has a mean duration of 12 days, minimum 9 days, maximum 15 days, based on a total of five observations. During this period the prothorax of the prepupa increases in size and becomes rounded and swollen while the lateral edges of the meso- and metathorax also become swollen and filled with fluid. A conspicuous occurrence during the prepupal stage is the appearance of the developing adult eye and the degeneration of the larval eye. The developing eyes appeared after one or two days in the prepupal stage as small pigmented spots just anterior to the prothoracic legsbeneath the prothoracic integument. Gradually the pigmented spots enlarged and appeared to migrate to the posterior end of the swollen prothorax, ventral to the prothoracic spiracle, where they remained until the prepupa moulted to the pupal stage. The pupa is decticous with the appendages clearly apparent and free from the body (fig. 3b). On each of the tergites of the third and fourth abdominal segments of the pupa are two pairs of minute hooks. Milliron (1940) observed median fleshy lobes bearing similar hooks on the second and third abdominal tergites of a mantispid pupal sheath. The function of the hooks in the chestnut mantispid is not known but it is likely that they may assist the pupa in emerging from the cocoon. The duration of the pupal stage was 20-28 days. The early pupa was a translucent creamy colour with prominent colourless eyeswhich had a light brown central portion. As the pupa aged, pigmentation of the eyes spread and by the eighth day the eye was black. On the 1lth or 12th day sclerotization of the thorax became apparent and the abdomen darkened two or three days later. Just prior to the emergence from the cocoon the pupa was a rich brown colour.

Adult emergence The pupa bites its way out of one end of the cocoon and moves around actively for approximately 45 minutes before anchoring itself firmly to the substrate by means of the legs. The pupal cuticle splits down the median dorsal line of the head and thoracic segments and is then pushed back by the body contractions of the emerging imago. First the head, then the legs were released from the exuviae and emergence was completed in about ten minutes. The wings were soft, white and crumpled and held above the abdomen of the imago at an angle of about 45" while the prothorax was bent dorsally. While the cuticle hardened the prothorax straightened and the wings were spread back over the abdomen. Finally the imago discharged a meconium from the anus. The adults were fed on Drosophila adults but copulation did not occur and no eggs were laid in captivity.

COCOON SPINNING 13EHA\'IOUR The spinneret in the third stage chestnut mantispid is located at the distal end of the abdomenand is themodified caudal sucker. The spinneret and ninth abdominal segment bear conspicuous posteriorly directed setae which seem to be tactile receptors relaying informarion about the substrate upon which the insect is spinning. The total cocoon spinning period was approximately 24-30 hours depending partly on the size Bissett and Moran : A South African mantispid 89 of the larva and the suitability of the substrate. If the larva constructs a cocoon on a flat open substrate, foundation laying takes longer than if the spinning larva were en- closed in a spider's cocoon or rounded hollow. When the cocoon is completecl ir is oloid and comprises a dense mass of white silk. The size of the cocoon varies considerably depending on the size of the larva when spinning commences. This is in agreement with the situation in the wasp Latra analis Fabricius (Smith 1935) where cocoons vary in qize according ro the size of the larvae. Cocoon spinning behaviour in the chestnut mantispid can be divided into three phases starting with an "initial phase" leading through an "intermediate phase" and ending in a "final phase" which is the dominant cocoon spinning period. The description below is based on observations of spinning in about 25 mantispid larvae.

The initial phase The initial phase is characteristically the period during which the foundation of the cocoon is built up. At the onset of spinning a thin stream of silk issued from the spinneret and by movement of the last five abdominal segments the silk was stuck to any convenient vantage points in the environment above, below or to the side of the larva. Every two or three minutes during this "haphazard" foundation laying the larva adjusted its position by three to six clear muscular contractioris which moved anteriorly along the body. With the dorsal surface of the prothorax and spinneret acting as points of support, these muscular movements altered the position of the larva and silk was thus laid down in different positions. In this way the larva constructed a network of silk around its body and the initial phase continued with the larva elaborating the silken network. This was achieved by the larva spinning over gaps in the network and frequently strengthening the framework by laying further silk over existing strands. Frequently just after the larva had changed its position in the developing cocoon it thrust its head outwards onto the silken framework and we suggest this may act as a tac~ilesignal initiating the intermediate phase; tactile receptors on the head may relay information to the animal about the extent and number of gaps in the existing framework. It is interesting to compare this with the statement by Jenkins (1958) who worked on the beetle Dianous coerulescens Gyllenhal which has a posterior spinneret, and who says : "Gaps in the framework [of the cocoon] seem to be checked by use of the antennae and mourh parts".

Thz itltermediate phase The intermediate spinning phase includes behaviour patterns characteristic of both the initial and final phases. In the initial phase the larva spins "haphazardly" to complete the foundation of the cocoon and movement in the developing cocoon was restricted to irregular small positional alterations accomplishrd by posterior to anterior niuscular contractions. In the final phase, movements of the body were regular, predictable and comprised rotations about the longitudinal and transverse axes. Silk was laid down in the final phase by "figure-of-eight" movements of the spinneret. In the intermediate phase, initial and final stage spinning characteristics were irregularly alternated depending upon the completeness of the surrounding cocoon network but the rotations in the cocoon in the intermediate phase were similar to those in the final phase. If the spinneret of the larva during spinning in the intermediate phase encountered an area of the cocoon in which there were gaps, the larva reverted temporarily to the initial haphazard spinning phase patching and strengthening gaps in the cocoon. If the spinneret passed over an area in the cocoon which was reasonably complete and devoid of 90 J. ent. Soc. sth. Afr. Vol. 30, No. 1, 1967

gaps the larva would proceed with "figure-of-eight" spinning and regular movements in the cocoon characteristic of the final phase. Thus the intermediate phase is very similar to the final phase but depending on the condition of the cocoon the animal can interrupt "figure-of-eight" spinning and revert to "haphazard" spinning characteristic of the initial phase. Eventually all gaps in the foundation of the cocoon are effectively covered in the intermediate phase and the dominant final phase commences. The jnal phase During the final phase, spinning is always achieved by "figure-of-eight" movements of the spinneret. "Figure-of-eight" cocoon-spinning movements have also been reported for the braconid Apanteles congregatus Say (Cushman, 1918) but in this case modified mouth parts form the spinneret. If a gap was artificially cut in the cocoon after the final stage had commenced "figure-of-eight" spinning continued uninterrupted. This distinguishes the final phase from the intermediate phase, for in the latter "figure- of-eight" spinning is interrupted when a gap in the cocoon is encountered. Essentially spinning behaviour in the final phase was characterized by regular rotations of the larva in the cocoon which interrupt the spinning, the movements com- prising seven rotations of the larva about the longitudinal axis and an eighth "reversal" movement in which the larva alternated the position of the head and spinneret in the cocoon anteriorly and posteriorly. These eight rotational movements of the larva in the cocoon were repeated approximately every 50 minutes and after about 25 such behavioural sequences the cocoon was complete. The sequence of events during final phase spinning can best be explained by reference to figs. 4a-k. After any rotational movement in the cocoon the larva was fully extended longitudinally and started "figure-of-eight" spinning in this position. Silk was always laid down on the surface of the cocoon opposite to the position occupied by the larva. The A A silken "figure-of-eight" became progressively largrr (fig. 4a) as the spinnerct moved towards the head and the larva became doubled UD. Eventuallv as the suinneret a~~roached A the head and after a predetermined time the animal extended its body again and three muscular contractions occurred, the first of which started at the posterior end ofthe body and ran anteriorly, while the second started anteriorly and ran posteriorly and the third was a replica of the first. By means of these contractions th~larva rotated through a predetermined anqle in the cocoon. Reversal movements were also accompanied by muscular contractions of the body, three to six of which passedfrom posterior to anterior before each reversal. After the reversal movement the same spinning sequence com-

EXPLANATION OF FIGURES Fig. 4. Diagrams 4a-k illustrate the final phase in the cocoon spinning sequence of the chestnut mantispid. The diagrams on the left in each row show the position of the third instar larva in the developing cocoon, dorsal surface of the larva stippled, the arrow indicating the direction of movement of the spinneret during '

ANTERIOR A NT E RIOR

DORSAL

L L L E E F H F VENTRAL 8 DORSAL F T T T T

POSTERIOR EO! POSTERIOR H d

P" ROTATION

4b REVERSAL

ROTATION 1 92 J. ent. Soc. sth. Afr. Vol. 30, "Vo. 1, 1967

ROTATION 2

ROTATION 3

4f ROTATION L,

4Y ROTATION 5 Bissett and Moran : A South African mantisbid 93

ROTATION 6

ROTATION 7

REVERSAL

4k ROTATION 3. ent. Soc. sth. Afr. Vol. 30, No. 1, 1967 prising seven rorarional movements was initiated and repeated (figs. 4c-4j). The angle through which the larva travelled during each of the seven rotational movements was predictable in all spinning sequences. Thus the first rotation after the reversal (figs. 4c-d) was through approximately 90°, the second (figs. 4d-e) through 90°, the third (figs. 4e-f) through 70°, the fourth (figs. 4f-g) 60°, the fifth (figs. 4g-h) 60°, the sixth (figs. 4h-i) 70°, and finally the seventh rotational movement (figs. 4i-j) through approximately 80'. The timing of the seven rotational movements in a spinning sequence was also constant in all spinning sequences. Immediately after the reversal (fig. 4c) the animal spun for approximately five minutes before rotating in the cocoon, after this rotation (fig. 4d) about seven minutes were spent spinning before the next rotation (fig. 4e), another seven minutes before the next rotation (fig. 4f), then seven minutes before the nzxt (fig. 4g). 79 minutes before the next (fig. 4h), then seven minutes (fig. 4i), six minutes (fig. 4j) and finally three minutes just before the reversal (fig. 4k). The areas of the cocoon covered by silk during each of the spinning periods which occur between rotations are illustrated diagrammatically in the right hand columns of figures 4a-k. It can be seen that after two spinning sequences the entire inner surface of the cocoon has been effectively lined with silk. During the final phase of cocoon spinning the sequence described above and depicted in fig. 4 is repeated about 25 times once every 50 minutes with predictable times spent spinning in each position between the reversal movements and rotations occurring through predictable angles until cocoon spinning is complete. In the insects generally cocoon spinning behaviour has only been fully reported for the larva of the moth Dictyoploca japonica Butler (Yagi, 1926) and the silk worms Bombyx mori L. (Yokoyama, 1951) and Platysamia cecropia L. (Van der Kloot & Williams, 1953 a and b, 1954). The latter accounts are detailed and deal with the behaviour of Platysamia in constructing a cocoon and also with the manner in which this behaviour may be artificially modified. The method of cocoon spinning in Platysamia however cannot be compared with that in the chestnut mantispid as the two animals are entirely different in this respect and consequently cornparisins are not attempted here. It is worth noting however that work on the cocoon spinning behaviour of silk worms is hampered by the fact that cocoon spinning is complex and that after the outer envelope has been spun observation of the spinning larva is almost entirely obscured. In the chestnut mantispid, on the other hand, cocoon spinning behaviour is simple, predictable and not significantly obscured as the silken envelope develops. Thus the chestnut mantispid offers an excellent opportunity for a study of the controlling neural mechanisms and environmental factors influencing this innate, cocoon spinning, behaviour pattern.

SUMMARY

1. The life history of an unidentified South African mantispid is described. 2. The eggs in this species are stalked, laid in batches and give rise to first stage active triungulin larvae. On finding the eggs of spiders the triungulin feeds under- going hypermetamorphosis which is complete by the end of the third instar. The mature third instar larva constructs a cocoon. 3. Cocoon spinning behaviour may be divided into an "initial phase", an "intermediate phase" and a dominant "final phase". The characteristics of these phases are fully described. 4. The development of the prepupa and pupa within the cocoon and the adult emergence is also described. Bissett and Moran : A South African mantispid

REFERENCES

CUSHMAN, R. A. 1918. Notes on the cocoon spinning habits of two species of braconids (Hym.) Proc. ent. ,Sot. Wash. 20: 133-6. ELTRINGHAM. H. 1932. On an extrusible glandular structure in the abdomen of Mantisba styriaca Poda. Trans. R. ent. Soc. ~ond.-80: 103-5. HUNGERFORD, H. B. 1939. A note on Mantispidae. Bull. Brooklyn ent. Soc. 34: 265. JENKINS, M. F. 1958. Cocoon building and the production of silk by the mature larva of Dianous coerulescens Gyllenhal (Coleoptera: Staphylinidae). Trans. R. ent. SOC.Lond. 110:--.. --287-501. . KASTON, B. ,J. 1938. Mantispidae parasitic on spider egg sacs. ,71. N. Y. ent. Soc. 46: 147-53. MAIN, H. 193 1. A preliminary note on Mantispa. Proc. R. ent. Soc. Lond. 6: 26. McKEOWN, K . C. & V. H. MINCHAM. 1948. The biology of an Australian mantispid (Mantispa z'ittata Guerin). Aust. 2001. 11: 207-24. MILLIRON, H. E. 1940. The emergence of a Neotropical mantispid from a spider egg sac. Ann. ent. Soc. i4m. 33: 357-60. SMITH, C. E. 1935. Larra analis Fabricius, a parasite of the mole cricket Gryllotalpa hexadac- tyla Perty. PTOC.ent. Soc. Wash. 37: 65-82. SMITH, R. C. 1934. Notes on the Neuroptera and Mecoptera of Kansas with keys for the identification of species. 3. Kans. ent. Soc. 7: 120-45.- . TJEDER, B. 1966. Personal communication. VAN DER KLOOT, MI. G. & C. M. WILLIAMS. 1953a. Cocoon construction by the Cecropia silkworm: I. The role of the external environment. Behaaiour 5: 141-56. 1953b. Cocoon construction by thr Cecropia silkworm: 11. The role of the internal enviroment. Behaoiour 5 : 157-74. 1954. Cocoon construction by the Cecropia silkworm. 111. The alteration of spin- behaviour by chemical and surgical techniques. Behaviour 6: 233-55. WERNER, F. G. & G. D. BUTLER. 1965. Some notes on the life history ofPleea- banksi (Neurop- teia: Mantispidae). Ann. ent. Soc. Am. 58: 66-8. YAGI, N. 1926. The cocooning behaviour of a Saturnian (Dictyoploca japonica). J. exp. 2001. 46: 245-59. YOKOYAMA, T. 1951. Studies on the cocoon formation of the silkworm, Bombyx mori. Bull. seric. Ex@. Stn. Japan 13: 183-260.

Manuscript received January 1 1, 1967 Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 1583

Reference Citation: Bissett, J. L.; Moran, V. C. 1967 [1967.??.??]. The life history and coccon spinning behaviour of a South African mantispid (Neuroptera: Mantispidae). Journal of the Entomological Society of Southern Africa 30:82-95.

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