
Results
The Larva.
In Plate 23, Fig. 4, is a sketch of the full grown living larva indicating the zones of colouring. The head and front thoracic region appears black, and forms the front dark stippled zone (BZ). This is due to the presence of blue and black pigment in what appear to be special cells below the integument. Reddish or grey flattened cells also are found here. The upper middle of the larva is chocolate colour (GZ), due to the presence of food in the mesenteron (ME), and especially to the two long brown gastric diverticula (D, D) which stretch down, often below the mid section of the animal. In the full grown larva these chocolate brown tubes are partially masked by four long bands of grey fat cells (GF). In the lower mid region of the body there is a pink zone (PZ), due to the pink granules in the much coiled malpighian tubules. The end section of the body is mainly hyaline and yellowish, and forms a clearly marked region (CZ).
Imaginal discs and gonads were found in one Waipu specimen only 19 mm in length. In this larva the granules in the malpighian tubes had already been cleared cut, and this part of the animal was hyaline. There are no imaginal discs in connection with the luminescent organs. The silk glands appeared empty, but were full length. Their fate in the adult has not yet been worked out, but will be studied if material is available.
On Pl. 25, Fig. 7, is a ventral view of the last segment. At (CO) is a ventral band of short, sharp comb-like setae arranged in groups. This band does not spread quite up to the dorsal surface and is the largest of several such bands which are indicated diagrammatically in Fig. 4A on Pl. 23. These more anterior bands of setae which are double, are not easy to see, but Fig. 4A gives the plan in one larva in which they were clearest. Above the posterior band shown in Pl. 25, Fig. 7, passing forwards, there is another band, then five areas still further forwards. These bands enable the larva to grip the silk and mucus runway in which it normally rests. The undersurface of the thorax is thrown into narrow cross bands which help the larva to pull its body forwards.
As is shown in Pl. 25, Fig. 7, there are longitudinal muscles arranged in three groups (LM) by the contraction of which the larva is able to pull forwards the entire segment in which the light organ lies. When pulled forward into this position, this segment lies within the seventh segment—that is, the one in front, and this telescoping tends to bring together blue pigmented cells and fat, which form a dense area. It might be assumed that the larva uses this power to douse its light. Observation of living larvae does not support this idea, though anatomically it seems quite possible. The normal position of the eighth segment in both luminescent and non-luminescent larvae is extended.
The Head.
In Pl. 25, Fig. 7B, is the upper surface of the head of a larva killed in Carnoy's fluid. The mandibles (J) are large and sharply toothed. Below at (ML) are what may be the bluntly toothed maxillae. In between, the basket-like structure (BB) becomes everted in penetrating fluids like Carnoy, but is not normally seen in dead specimens. This basket has an aperture in front, and it appears to be an epipharynx, but its nature is not properly understood. Apparently the food is gripped by the mandibles and sawed into suitable lengths by the maxillary teeth (see Pl. 24, Fig. 6B). At each side of the head is the base of the antenna (A), and behind this is an eye (E2) which appears to be made up of a single facet but more than one ommatidium, and is therefore strictly an ocellus. Below this eye, in line with the bottom of the antenna, at (E) is another peculiar pigmented sense organ which more closely resembles a true lateral ocellus Like most startled nocturnal

insects, the larvae do not move at first when a flashlight is directed at them: this does not say they lack light perception; but their eyes probably only function as sense organs suitable to let them know when night has fallen, and dawn has come. The glow-worms do not see as well as the larvae of the tiger beetle, which live in holes in clay banks. The glow-worm depends on its snare, the tiger beetle larva on its sight and quickness.
The Gut of the Larva.
In Pl. 24, Figs. 5 and 6, are the parts of the gut, as seen when the last segment is cut off and the viscera pulled out by the head. The oesophagus (O) is markedly contractile. The chocolate coloured diverticula (D) show muscular movements, not apparently peristaltic, but such as would be produced by contraction of scanty circular muscles. No doubt it is by such contractions that the mucous droplets are expelled at intervals from the mouth, as the snare is being constructed. Owing to the position of emergence of the diverticula from the oesophageal valve, the latter probably controls the issue of mucus. The oesophageal valve is yellow in colour.
The mesenteron was never found quite empty in newly caught specimens, but if the larvae are starved for several days, the gut contents move down leaving the upper part empty. Food was never found in the intestine (IN) except on one occasion when the digested material was observed in the lower part of the mesenteron; it quickly passed through into the intestine and was immediately voided. The food contents seen in the mesenteron always appeared brownish. Recently a larva was seen to regurgitate a drop of food. In Pl. 24, Fig. 5, the four parts of the malpighian tubes (1–4) are shown. The first part (1) is yellowish and hyaline, the second and longest part (2) is of a larger bore, very opaque, and has a pink colour in direct light. The third part (3) very gradually passes into a thinner and more transparent region of part (4); this region is usually disposed in a zigzag fashion, covered with thin connective tissue, and provides the necessary extensile region needed as the animal contracts and lengthens quickly as it moves along; part (4) is the swollen luminescent organ, which, like the lowest part of part (3), is transparent and yellowish. The reflector is glistening white, and when cleared of attachments rapidly floats to the top of the water in the dissecting dish, resembling a tiny boat. These regions of the gut are depicted at a higher magnification in Pl. 24, Fig. 6. When the tubule is pulled out from the living animal, one occasionally sees some of the granules from part (2) passing into the lumen of part (1). It is this process which, before pupation, leads to the adult malpighian tubes being cleared of all the excretion granules collected during larval life. The adult's malpighian tubes are like part (1) of the larval tubes. Part (1), of course, leads into the gut proper. The urine (?) granules in part (2) gradually become less and less towards the distal end, until the cells become hyaline Cross sections of the region (3) reveal that the cells do not always occupy the complete area of the tubule, one side here and there showing a gap, as in Pl. 24, Fig. 6A. This is also shown in Pl. 1, Fig. 5 (C), of the paper by Gatenby & Ganguly (1958). The significance of this method of packing the cells is not known At the region at part (3A) in Pl. 24, Fig. 6, the lumen is still patent, but it appears to be shut off at (LN) by an area of about three cells in length. No granules were ever seen inside the lumen of the living luminescent organ.
In Pl. 24, Figs. 5 and 6, at CT, the connective tissue binding of part (3) and (3A) is drawn in diagrammatically to show that this region is so completely bound that the intestine and the four lower parts of part (3) cannot usually be separated without breaking. In Pl. 24, Fig. 5, the proportionate length of the meanderings of part (3) are not fully shown. A less diagrammatic drawing of this region in an extended animal is shown in Pl. 25, Fig. 7 (CT and M).

The silk glands extend far down to the mid region, and then each one turns up at its lower part (SI) in Pl. 24, Fig. 5. Fat cells (F) are closely bound to the silk glands.
Tracheal System.
In Pl. 23, Fig. 1, is a semi-diagrammatic plan of the tracheal trunks. There are two main tubes (L, L) passing from a fork which goes to head and brain, right down to the reflector and luminescent organ (LO). On the way down these give off branches (T, T) to the viscera. The main trunk leaving the reflector (LO) branches forward (lower T). At intervals the main trunk receives tubules (B) which have branched under the integument at each site. There are no tracheal openings to the exterior, the system being apneustic.
In the pupa, Pl. 23, Fig. 2, there are at least six tracheal openings, each in the mid region of the segment. A higher power drawing of one of those is given in Pl. 23, Fig. 3, the opening at TA, longitudinal trunk at L.
The tracheal system of the imago has not been studied but is probably normal for the family.
The Luminescent Organ.
In Pl. 27, Fig. 12, is a good photograph of the light organ (LO). There are four tubes seen in transverse section in Pl. 26, Fig. 9. In this region the reflector (R) lies underneath, and the tubes are not bound to this, but lie in depressions in the reflector. The intestine and rectum (IN) also lie free, but as far as can be ascertained, the light organ itself is not free to move up or down according to the position of the connectives (CT), and the contraction and relaxation of the longitudinal muscles (LM). The light organ is shown well at (U) in Pl. 27, Fig. 10, the lumen more clearly at (LO) in Pl. 27, Fig. 12. Although the light given by small larvae appears to be bright, their luminescent organ is comparatively small. In Text-fig. 4 (A), is an outline of the adult larval organ, in (B) that of a larva 5 mm in length.
The living cells of the light organ are hyaline and free from granulation except for mitochondria packed in the cytoplasm of the cell.
Lengths of Parts of the Viscus of the Larvae, Etc.
Oesophageal valve, 1.25 mm; silk glands, 22 mm; mesenteron, 10 mm; gut diverticulum, 12 mm; malpighian tube, yellow region (part 1), 2.5 mm; pink region (part 2), 9 mm; (part 3A, difficult to measure); luminescent organ, 0.9 mm; altogether about 15 or 16 mm. This, from typical larvae, but the lengths in some cases obviously depend on the amount of stretching and size of larvae.
The writer's drawings had to be made mostly freehand, as no camera lucida was available at the time. The average larva was about one inch (25 mm) long. One from Waipu, apparently a male, nearly ready to pupate, that is with imaginal discs and visible gonads, was only ¾ inch (19 mm), yet another from the same cave was 1 ¼ inches (about 30 mm) long, and it had no visible imaginal discs in whole mount.
Nervous System.
The nervous system consists of the larger cerebral ganglia (brain), para-oesophageal commissures and sub-oesophageal ganglia, the usual double ventral cord, three ganglia in the three thoracic segments, and a ganglion just behind each abdominal segment, except in that segment containing the light organs. The brain lies just behind the head capsule, not inside it (Pl. 25, Fig. 7B, BR). In Pl. 25, Fig. 7A, are the last three segments, and the supply to the region containing the light organs (LO) comes from the ganglion (NA) in the penultimate segment. This ganglion sends down a nerve which soon branches, one part to the muscles, the other continuing down to the region of the light organs and anal papillae. One

branch (N) certainly comes from the scolophore organs, but it is difficult to follow the paths of the other small branches, some of which go to muscles. This means that it is not presently possible to say whether the central and the sympathetic visceral systems are brought down into the posterior part of the malpighian tubules which are the light organs, and to the reflector. In any case a cut anywhere below the letters (NA) in Pl. 25, Fig. 7A, will isolate the central nervous system and its anterior sense organs, from the light organs. Such a cut will also isolate the posterior part of the haemocoele from any possible hormonic influence arising in front parts of the nervous system, but it will shut off the oxygen supply arising from the segmental tracheole branches beneath the integument. The long tubular heart passes back to just near the border of the last segment. Thus the cut will also interrupt the heart, but will leave a portion of variable length according to the level of the cut below the ganglion (NA) in Fig. 7A. With these difficulties in view a few experiments were made.
Experiments on the Control of Luminescence.
On Pl. 23, Fig. 4, the arrows X, Y and Z show where a cut was made or a nylon thread was tied. The thread was Size A, white, Coats and Clark's, was fine enough for the experiments, and is very strong. Three main types of experiments were tried.
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(1) A tie or cut behind the head, isolating the head from the rest of the body. Head cut off in all cases.
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(2) A tie or cut between X and Y, usually in front of Y, body cut off.
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(3) A tie or cut at Z, isolating any nerve ganglia from the light organ, the body being removed.
It was very difficult to get a thread on to a struggling lighted up larva, before it doused. In (1) and (2), the larva could douse. In (3) the piece below Z, could not douse, and if the experiment was carried out on a doused larva, the luminescent organ lighted up immediately. If (2) was tried on a doused larva, it did not light up, but if (3) was then carried out, it lighted up If the lighted up piece (teased or not) from experiment (3) be covered with blood from a doused larva, it still continues to light up.
From these experiments it appears that the light is kept doused by the posterior ganglion in segment 7. In all cases the light does not turn off or come on suddenly, from which it may be concluded that some other organ is implicated, but not a humoral effect or hormone in the haemocoele fluid. It is likely, but there is no evidence, that the ganglion in segment 7, in some way prevents air from the tracheal reflector from diffusing to the light organ.
Before any conclusion can be drawn, the nerve supply to the light organs and tracheal basket must be examined, but suitable gold or silver preparations have not been made.
In the case of those glow-worms living outside on banks, at present it seems likely that the approach of evening is registered by the eyes of the larva, and a stimulus is sent down from the brain via the ventral nerve cord to the posterior ganglion, which then lifts the block which hitherto has kept air from diffusing across the small gap between the reflector and the light organs. The blood containing air from the reflector could be removed by contraction of the last segment, the swelling or contraction of which is entirely under the control of the larva. Note that in Pl. 23, Fig. 4, this segment is largely contracted, while in Pl. 25, Fig. 7, it is suffused with blood. The writer feels that this is in some way concerned with dousing.
Many attempts were made to dissect the posterior ganglion so that the nerve fibres could be followed to their end organs, but the cuticle was so tough that the task was beyond the skill and patience of the writer The question of the use of the eyes for the perception of night and therefore of lighting-up, was not solved.

Fig. 1—Plan of the larval tracheal system. Fig. 2—Exuvia of male pupa, ventral surface. Fig. 2A—Head and thora,x ditto, from side. Fig. 3—Tracheal trunk and opening (spiracle) of pupa. Fig. 4—Nearly full-grown larva, showing zones of colouration. The spread of lines between (CO) and (LO) represents light. Fig. 4A—Plan of banded rough areas for adhesion by combs.

Fig. 7—Last segment of larva from below. The reflector is black. Compare with Fig. 9. Fig. 7A—The three posterior segments of the larva showing position of the last abdominal ganglion (NA). Fig. 7B—Head of larva from above showing toothed mandibles (J) and serrated maxillae (ML) with everted basket (B) lying in between. Antenna base (A) and two pigmented eyes (E, E2).

Larvae are easily anaesthetized by being immersed in water containing the anaesthetic, but chloroform-water eventually killed them. It should be possible to find a more suitable anaesthetic, to dry the larvae, and then to paint over their eyes. It is to be expected that larvae so blinded will luminesce continually.
