ToxICOIf. Vol. 31. No.7. pp. 881-887. 1993. 4 ~ 9 3 S6.00 + .

00
PrUned in Great Britain. o 1993 PerpmonPress ltd
DELIVERY OFDUVERNOY'SSECRETIONINTO PREY BY THE
BROWNTREESNAKE, BOIGA IRREGULARIS
(SERPENTES:COLUBRIDAE)
WILLIAM K. HAYES,
1
PABLO LAviN-MURCI0
2
and KENNETII V. KARDONG
2
*
'Biology Department,Southern College Collegedale, TN 37315-0370, U.S.A.; and 2Department ofZoology,
Washington State University Pullman. WA99164-4236, U.S.A.
(Received23 September 1992; accepted II January 1993)
W. K. HAYES, P. LAvfN-MURCIOandK. V. KARDONG. DeliveryofDuvernoy's
secretion into prey by the brown tree snake, Boiga irregularis
(Serpentes:Colubridae). Toxicon 31, 881-887, 1993.-Manycolubridsnakes,
like themore venomouselapid andviperid snakes, can produce andinject an
oral secretion that is toxic and may present a human health risk. However,
colubrid oral toxins are produced in a Duvernoy's gland and delivered not
through a hollow fang, but instead by long, often grooved teeth under low
pressure. The possible role ofDuvernoy's secretion in functions other than
rapid killing ofpreymakeit importantto know howandwhere this secretion
is deliveredduringa feeding strike. We used ELISAanalysis todetermine the
quantity and proportionaldistributionofDuvernoy's secretion delivered into
theintegumentcomparedto thevisceraduringafeedingstrikebythecolubrid
snake Boiga irregularis. We detennined that only about54% (1-5mg) ofthe
secretion actually reached the viscera and that the rest remained in the
integument. Theamountreaching theviscera is about three toeight times the
i.p. LD
so
formice, butthesesnakesdependmoreonconstriction thantoxins to
killtheirprey.Consequently,deliveryofDuvemoy'ssecretionbyB. irregularis
is hypothesized to be partofa digestivefunction andits toxic propertiesa by-
productofthis role.
INTRODUcrION
SoME colubrid snakes produce oral secretions that exhibit mild (VEST, 1981a.b) to even
alarming toxicity (McKINSTRY, 1983; SAKAI et al., 1984), with reports ofhuman deaths
following bitesbysomeofthesecolubridspecies (MITTLEMAN andGoRIS, 1974; MINTON,
1990). Yetcolubridsnakeslacka venomgland. Insteadmanypossessa Duvernoy'sgland
(TAUB, : 1966, 1967), evolutionarily' homologous to the venom glands of elapids and
viperids (KOCHVA, 1965; KOCHVA and WOLLBERG 1970), butsurprisingly distinct from a
true venom gland. For example, Duvernoy's gland does not empty into a hollow fang.
Instead,itsductopensintotheoralcavitynearposteriormaxillaryteeth thatmayormay
not be grooved. Posterior maxillary teeth are often engaged during prey capture and
swallowing (e.g. KAROONG, 1980, 1986). But an open-grooved or non-grooved tooth
Author to whom correspondenceshould beaddressed.
881
882
W. K. HAYES tl 01.
offers no protected, enclosed channel passing through the integument and into the prey.
Consequently, the pliable skin of the prey can obstruct the tooth or its groove, making it
less effective as an instrument for introduction of secretion from oral glands (KARDONG
and YOUNG, 1991). Further, such teeth of colubrids cannot hold a pressure head generated
by Duvernoy's gland in the same way that the hollow fang of elapids and viperids allows
high-pressure delivery of a shot of venom (KARDONG and LAViN-MuRCIO, 1993). Finally,
Duvernoy's secretion may tranquilize prey or even lead to a protracted death. However, if
surgically deprived of secretion from Duvernoy's gland during prey capture, some snakes
nevertheless can and do kill prey by constriction as rapidly as when the secretion is
available (ROCHELLE and KARDONG, 1993). For these reasons and for reasons related to
the evolution of the venom apparatus, it has been argued that the function of the secretion
from Duvernoy's gland should be evaluated separately from the venom of elapids and
viperids (KARDONG, 1982a). To clarify the role of secretion from Duvernoy's gland in
feeding and to assess its medical implications in cases of human envenoming, the question
also arises as to just how much Duvernoy's secretion actually enters the prey. Our purpose
then was to detennine the quantity of Duvernoy's secretion delivered during a feeding
strike, and to compare how much remained in the integument with how much actually
reached the viscera.
We used the colubrid species Boiga irreguJaris, the brown tree snake. Because of its
ecological and economic impact (FRrrrs, 1988), the brown tree snake has received special
attention, giving us information on its basic feeding ecology (SAVIDGE, 1986, 1987, 1988).
Recent studies of the secretion of Boiga (Vest et a/., 1991; Weinstein et al., 1991) and the
morphology of its venom apparatus (Zalisko and Kardong, 1992; Kardong and Young,
1991) provide further details which relate to the quantity of venom released during
feeding.
MATERIALS AND METHODS
Subjects
Nine adult brown tree snakes (122-184cm in snout-vent length), collected on the island of Guam. were
maintained in glass aquaria in a room with temperature (28
C
C 2) and humidity (70%) controlled. The captive
snakes were fed every 2 weeks on laboratory mice (Mus musculw) for 9 months prior to experiments.
Experiment I
Each snake was permitted to strike and constrict an adult Swiss-Webster mouse (20.1-24.0 g) introduced into
its home cage. Their feeding behavior was recorded until the mouse appeared to be dead (exhibited no reflexes).
Then
9
the rodent was quickly retrieved, placed into a plastic bag, and frozen ( - 20e); another mouse sacrificed
earlier was offered to the snake for consumption.
The quantity of Duvernoy's secretion expended (dry mass) on each prey animal was measured to the nearest
milJigram by enzyme-linked immunosorbent assay (ELISA) of whole-mouse homogenates. After thawing, the
mice were homogenized 3-4 min in a 14-speed Osterizer blender with 150 ml buffer (0.05% Tween-20 in
phosphate-buffered saline (PBS); see HAYES et 01., 1992). Several 1.5 mJ aliquots of homogenate were centrifuged
5 min in an IEC Oinical Centrifuge. The supernatant was frozen at - 20
0
e until ELISAs were performed.
The ELISA methods are described elsewhere in detail (HAYES et al., 1992). Brieftyt venom extracted by
micropipet (e.g. VEST,1981b), but without aspiration from the Boiga specimens (4 months prior to study). was
injected into two rabbits using the modified method of induced granuloma. This method (HILUM et al'
9
1914)
takes advantage of a granuloma fonned within a perforated. plastic golf ball surgically implanted under the skin
of a rabbit. Fifteen days post-implementation, 200 pg of Duvernoy's secretion (antigen) in I mI PBS was injected
directly into the lumen of the implanted ball; 2 weeks later a booster injection of another 200 pg was injected into
the ball. Twenty-one days after tlJe booster, up to 21 ml of clear. yeJlow fluid was directly withdrawn from the
lumen of the ball using a two needle system. A sterile 22 gauge filter needle allowed air to pass into the lumen of
the ball as ftuid was being withdrawn through a second 22 gauge needle attached to a syringe. Polyclonal
antibodies (IgG) were isolated from the hyperimmune fluid by protein A-Sepharose affinity chromatography. A
883 Brown Tree Snake Bite
portion of the immune IgG was conjugated to horseradish peroxidase (HPO: Sigma type VI. P 8375. RZ = 2.9)
by the pcriodate method. After evaluation by ELISA. purification of the conjugate was deemed unnecessary.
Triplicate samples of homogenate from the nine struck mice were assayed on a single Dynatech Immulon I
microtitre plate. The plate also contained triplicate samples of homogenate from nine control mice (20-24 g)
injected with 0-4 mg venom in 0.5 mg increments. The venom. dissolved in 0.25-1.0 ml PBS, was delivered via
tuberculin syringe (25 g needle) just below the skin of the right mid-dorsal region. These control mice. which were
nccessary for preparing a standard curve. were treated otherwise in an identical manner as the experimental mice.
To assay venom in the mouse homogenates. wells of the plate were coated overnight at 4C with IgG in a
10 pg/ml PBS solution (99 pi/well). The wells were emptied (by slapping plate onto paper towels) and washed
three times with Tween-20/PBS buffer (130 pl;well). Next (since a blocking step was found to be unnecessary: cf.
HAYES et al., 1992), the samples of mouse homogenate (I : 1000 dilution in PBS) were added (99 pl,:well) and
incubated for I hr at room temperature. After the wells were emptied and washed as before, IgG-HPO conjugate
was added (99 pi/well) and incubated another hour at room temperature. Finally, after wells were emptied and
washed again, substrate solution [0.1 mt of 0.50/0 hydrogen peroxide added to 9.9 ml of 5-AS substrate (from
Sigma) in I mg/ml 0.1 M sodium phosphate/EDTA buffer, pH 5.95] was added (99 pi/well). As the plate was
agitated manually during the next hour, color development (absorbance) was read by an automatic microplate
reader (Titertek Multiscan MCC/340) at 450 and 620 nm (dual wavelength mode).
To develop a standard curve, absorbance values for control homogenates were plotted against the concentra-
tion of venom and a regression equation was fitted using the program Quattro Pro (version 3.0). The regression
equation was used to estimate the mass of venom expended (nearest mg) for each mouse based on the mean
absorbance of the three homogenate samples from each carcass.
Experiment 2
Three months later, seven snakes (of the nine from experiment 1) were observed as before as each struck and
constricted an adult male mouse (17-24 g) in its home cage. Immediately after death, the mouse was removed
from the snake's mouth, stored within a plastic bag, and frozen at - 20
G
e. Two months later, the mice were
thawed, quickly and carefuJIy skinned, and skin and viscera (skeleton, musculature, internal organs) placed in
separate bags. As in experiment I, each was homogenized with 150 m1 wash buffer, centrifuged, and the
supernatant frozen until ready for ELISA. Then, triplicate samples from three sets of homogenate (control,
viscera, skin) were assayed on a single microtitre plate to detennine concentrations of venom in both the viscera
and the skin.
Validity of ELISA
The coefficients of determination (r) for our regression equations indicate the amount of variation in
absorbance explained by the independent variable, quantity of venom present. The coefficients for the two
experiments were, respectively, 0.81 and 0.84 (explaining more than 800/0 of the variance). which indicated that
absorbance was a good indicator of the Duvernoy's secretion present in the samples.
Since the snakes deposited considerable quantities of saliva on the mice, we were concerned that our assay
might have measured saliva in addition to venom. To confirm that the antibodies were not cross-reactive with
Boiga saliva, we obtained saliva samples via microAipets as before from the oral epithelium of three of the live
snakes. These samples were lyophilized. weighed, reconstituted in PBS. and assayed together with Duvernoy's
secretion and venom from the midget-faded rattlesnake. Crotalus viridis concolor (as another control). In this
ELISA. the wells were coated and washed as before. Duplicate samples of saliva and venom (I mg/ml PBS) were
then added and serially diluted in ten-fold dilutions (I : 10) of PBS to 0.1 ng/mt. The wells were incubated 1.5 hr
at room temperature and then washed. IgG-peroxidase conjugate was added next and incubated 2 hr at room
temperature. After washing, substrate solution was added and color was read as before.
Detectability, a useful indicator of cross-reactivity, is defined as the minimum concentration of venom (10")
that can be detccted by the ELISA (MINTON el al., 1984; KAISER. el al., 1986; BoBER et al., 1988). The level of
detectability of Boiga Duvernoy's secretion was OJ)OI pgfml. Detectability for Boiga saliva was 1000 JJg/mI. a
difference of six orders of magnitude. C. v. concolor venom was not detectable. Thus, the antibodies appeared to
be very specific for Duvernoy's secretion.
RESULTS AND DISCUSSION
The quantities of Duvernoy's secretion expended by brown tree snakes when striking
mice and the survival times of prey are summarized in Table 1. The snakes expended
similar total quantities of venom in the two (means of 3.6 mg and 3.3 mg,
respectively). These amounts of Duvernoy's secretion are less than but within an order of
magnitude of those amounts of venom injected into mice by elapid and viperid snakes of
884 w. K. HAYES et al.
TABLE 1. DRY MASS OF DUVUNOY'S SECREIlON (VENOM) EXPENDED BY TIlE BROWN TREE SNAKE (Boiga irregularu)
DURING FEEDING AND TIME TO DEATH OF THEIR PREY, LIVE ADULT MICE (Mus museu/us)
Experiment 1 (n = 9) Experiment 2 (n = 7)
Dependent measures Mean S.E. Range Mean S.E. Range
Total venom expended (mg) 3.6 0.3 2-5 3.3 0.1 1.4-4.7
In viscera 1.8 0.2 1.0-2.7
In skin 1.5 0.2 0.5-2.2
Time to prey death (sec) 113 48 37-484 65 7.8 36-101
comparable size (HAYES et aI., 1992). However, the flow of Duvernoy's secretion in
colubrid snakes is very slow compared to venom release by elapid and viperid snakes
(CmszAR etal., 1992; HAYES, 1992; KAROONG and LAvlN-MURCIO, 1993), and may have
been continuous during the several minutes that the teeth engaged prey tissues.
Considering the minimum of 6.7 mg venom available in the Duvernoy's glands of adult
B. irregularis(WEINSTEIN etal., 1991; CmszAR etal., 1992; VEST etal., 1991), the snakes of
the first experiment expended approximately 54.5% of their available venom by the time
their prey was dead. This value is equal to the highest percentage of available venom
expended by any elapid or viperid snake for which comparable data are available (HAYES
et al., 1992); however, the venom is not released instantaneously as in other venomous
snakes. An additional quantity of venom would likely have been secreted during the
swallowing process had we permitted the snakes to do so. Unlike the venom glands of
most elapids and viperids (KocHVA, 1978), the Duvernoy's gland of B. irregularislacks an
extensive extracellular storage reservoir (ZALISKO and KARooNG, 1992). Therefore, this
high percentage expended is a likely consequence of the low volume of ready venom
within the lumen of the gland.
Results of experiment 2 indicate that approximately 54.5% of the total venom released
was delivered into the viscera of prey, while 45.50/0 remained embedded in the skin (Table
I). The fact that total venom expenditure did not differ significantly between the two
experiments gives us confidence that the separation of mouse skin from viscera in the
second experiment did not confound our results. The value of 54.5% for envenoming
efficiency is considerably below that of elapid and viperid snakes (MORRISON etal., 1982,
1983; HAYES et al., 1992). However, prior studies of envenoming considered only that
venom on the surface of the integument which could be washed from it; these values for
Boiga include venom embedded in the skin of prey, so comparisons must be treated
cautiously.
The mean of 3.6 mg venom expended (experiment 1) when adjusted for mice averaging
21 g represents a dose of 171.4 mg/kg. The Lp. LD
50
for B. irregularis is reported to vary
between 10.5 and 34.1 mg/kg (VEST et al., 1991; WEINSTEIN et al., 1991, 1993). Thus, the
amount of Duvernoy's secretion delivered during a feeding strike is well above the lethal
dose (50%) necessary to kill mice. If only the portion of the secretion that entered the
viscera (experiment 2) is considered (1.8 mg 0.2), this still gives an adjusted value of
904 mg/kg, three to eight times the lethal dose (500/0) depending upon which value is
compared (34.1 or 10.5 mg/kg).
This success in getting a lethal dose of DuvernoY,s secretion through the integument
and into the viscera is particularly noteworthy for two reasons. First, delivery of
Duvernoy's secretion during prey capture does not significantly contribute to rapid death
of the prey. Where prey death was compared for B. irreguJaris with and without use of
885 Brown Tree Snake Bite
Duvemoy's secretion, it was found that no difference occurred in the rate of prey death
(ROCHELLE and KARDONG, 1993). The brown tree snake uses jaw pressure (ROCHELLE and
KARDONG, 1993) and most prominently constriction (CHISZAR, 1991; ROCHELLE and
KARDONG, 1993) to kill prey quickly. On occasions where prey escape or are removed
before constriction brings death, they may become sluggish and even eventually die after
many minutes or several hours (McCarn, personal communication). This suggests that
Duvernoy's secretion can tranquilize prey or even produce a protracted death. However,
there is no published evidence that Boiga Duvernoy's secretion alone can equal the rapid
prey death characteristic of the true venoms used during feeding by elapid and viperid
snakes.
Second, the posterior maxillary teeth of Boiga, associated with the single duct from
Duvernoy"s gland (ZALISKO and KARDONG, 1992), bear an open groove. Thus, unlike the
hollow teeth of elapids and viperids that form an enclosed venom channel within the fang,
these grooved teeth of Boiga are an open channel and cannot sustain a pressure head to
deliver a pulse of secretion deep into tissues (KARDONG and LAViN-MuRCIO, 1993). Our
results suggest that as a consequence, a significant portion (45.5%) of the delivered
secretion remains lodged in the integument until the prey is dead.
Taken together, these results support the view that Duvernoy's secretion in most
colubrids does not playa primary role in rapidly killing prey as do the venoms of elapids
and viperids (KARDONG, 1982b). Further, the biological role of Duvernoy's secretions in
most colubrids is most likely related to some other primary function, such as digestion
(KARDONG, 1982a). The presence of proteolytic enzymes in Duvernoy's secretion from
Boiga (WEINSTEIN et aI., 1992) supports this suggestion. The experimental evidence in
other colubrid species of a digestion-promoting effect of Duvernoy's secretion (JANSEN,
1983; RODRiGUEZ-RoBLES and THOMAS, 1992) further supports this hypothesis. Thus,
toxicity of Boiga Duvernoy's secretion may be a simple by-product of this primary
digestive role.
The digestive action of Duvernoy's secretion can act directly upon prey tissues to
promote chemical digestion. We hypothesize that Duvernoy's secretion may additionally
serve to open holes chemically in the tough integument of the swallowed prey to pennit
later entry of digestive enzymes released by the snake's digestive tract. Although we
removed mice before swallowing began, the location of the grooved teeth results in their
regular engagement with the integument of the prey as the jaws ~ w l k over it, as in other
colubrids (KARDONG, 1986). Consequently, during swallowing cycles, further quantities of
Duvernoy's secretion from the gland could be introduced into the skin. Because the prey is
dead when such swallowing begins, no active spread of the secretion by the circulatory
system would occur, and for the most part the secretion would remain at the sites in the
integument where the teeth initially deposited it.
If one biological role for Duvernoy's secretion is to open chemically the integument of
the prey for entry of digestive tract enzymes, then this would help to account for why
during prey capture such a large portion (45.5%) of the secretion remained embedded in
the skin and why even further secretion is likely introduced during swallowing.
Acknowledgements-Very special thanks go to Dr NANCY MAGNUSON who generously contributed her time and
laboratory resoures during the running of ELISA analyses. The School of Veterinary Medicine kindly loaned us
use of an ELISA plate reader, J. FOLWELL helped with preparation of solutions, and M. ROCHELLE helped with
husbandry of snakes. Supported in part by U.S. Fish & Wildlife Service and NSF (Animal Behavior,
BNS-8820091), and by CONACYT (Mexico).
886
W. K. HAYES et al.
REFERENCES
BoBER. M. A. GLENN. J. L.. STRAIGHT. R. C. and OWNBY. C. L. (1988) Detection of myotoxin proteins in
various snake venoms. Toxicon 26, 66>-693.
CHISZAR. D. (1991) The behavior of the brown tree snake: A study in applied comparative psychology. In:
ComemporarJ'lssue in Comparative Psychology, pp. 101-123 (DEWSBl:RY. D. A.. Ed.). Sunderland: Sinauer
Assoc.
CHlSZAR, D., WEINSTEIN, S. A. and SMITII. H. M. (1992) Liquid and dry venom yields from brown tree snakes,
Boiga irregularis (Merrem). In: Contributions in Herpetology, pp. 11-13 (STRIMPLE, P. D. and STRIMPLE, J. L.,
Eds). Cincinnati: Greater Cincinnati Herpetological Society.
FRITTS. T. H. (1988) The brown tree snake, Boiga irregularis. a threat to Pacific islands. U.S. Fish Wildl Serv.,
Bio/. Rep. 88(3), 36 pp.
HAYES. W. K. (1992) Factors associated with the mass of venom expended by prairie rattlesnakes (Crotalus v.
.,iridis) feeding on mice. Toxicon JO, 449-460.
HAYES. W. K. KAISER. I. I. and DUVALL. D. (1992) The mass of venom expended by prairie rattlesnakes when
feeding on rodent prey. In: Biology of Pit.,ipers, pp. 383-388 J. C. and BRODIE, E. D. JR, Eds).
Tyler. TX: Selva.
HILLAM. R. P. L., TENGEllDY. R. T. and BROWN, G. L. (1974) Local antibody production against the murine
toxin of Yersinia pestis in a golf granuloma. Infect. Imm. 10, 458-463.
JANSEN, D. W. (1983) A possible function of the secretion of Duvernoy's gland. Copeia 1983, 262-264.
KAISER. I. I.. MIDDLEBROOK. J. L.. CRUMRINE. M. H. and STEVENSON, W. W. (1986) Crossreactivity and
neutralization by rabbit antisera raised against crotoxin. its subunits and two related toxins. Toxicon 24,

KARooNG, K. V. (1980) Evolutionary patterns in advanced snakes. Amer. Zool. 20, 269-282.
KARDONG. K. V. (1982a) The evolution of the venom apparatus in snakes from colubrids to viperids and elapids.
Mem. Inst. Butantan 46, 10>'-118.
KAROONG, K. V. (1982b) Comparative study of changes in prey capture behavior in the cottonmouth
(Agkistrodon piscivorus) and Egyptian cobra (Naja haje). Copeia 1982, 337-343.
KARDONG, K. V. (1986) Kinematics of swallowing in the yellow rat Elaphe obsoleta quadrivitlata: a
reappraisal. Jpn. J. HerjJ. 11, 96-109.
KARooNG, K. V. and P. A. (1993) Venom delivery of snakes as and
systems. Copeia 1993, 644-650.
KARDONG, K. V. and YOUNG. B. A. (1991) Fangs and snakes: how do open grooves inject venom into enclosed
spaces? Amer. Zool. 51A. _
KocHVA. E. (1965) The development of the venom gland in the opisthoglyph snake Telescopus fa/lax: with
remarks on Thamnophis sirtalis (Colubridae. Reptilia). Copeia 1965, 147-154.
KOCHVA, E. (1978) Oral glands of the Reptilia. In: Biology of the Reptilia, Vol. 8, pp. 43-161 (GANS. C. and
GAN5, K. A., Eds). New York: Academic Press.
KOCHVA. E. and WOLLBERG. M. (1970) The salivary glands of Aparallactinae (Colubridae) and the venom glands
of laps (Elapidae) in relation to the taxonomic status of the genus. J. Linn. Soc. (Zool.) 49, 217-224.
McKINSTRY. D. M. (1983) Morphologic evidence of toxic saliva in colubrid snakes. A checklist of world genera.
Herp. Rev. 14, 12-15.
S. A. (1990) Venomous bites by nonvenomous snakes: an annotated bibliography of colubrid
envenomation. J. wi/d. Med. 1, 119-127.
S. A. WEINSmN. S. A. and WILDE III, C. E. (1984) An immunoassay for detection of
North American pitviper venoms. Clin. Toxic. 22, 303-316.
MIlTLEMAN. M. B. and GoRIS, R. C. (1974) Envenomation from the bite of the Japanese colubrid snake
RMbdophis tigrinus (Boie).. Herpetologica JO, 113-119.
MORRlSON, J. J., PEAllN. J. H. and A. R. (1982) The mass of venom injected by two Elapidae: the
taipan (OxyuranUs sculelJatus) and the Australian tiger snake (Notechis scuta/us). Toxicon 20, 739-745.
MORJUSON, J. J. PEAJlN. J. H., CHARLES N. T. arid COULTER, A. R. (1983) Further studies on the mass of venom
injected by elapid snakes. Toxicon 21, 279-284.
ROCHELLE, M. and KARooNG, K. V. (1993) Constriction vs. envenomation in prey capture by the brown tree
snake Boiga irregu/aris (Squamata: Colubridae). Herpetologica. 49, 297-300.
RODlUGUEZ-RoBLES, J. A. and THOMAS, R. (1992) Venom function in the Puerto Rican racer. Alsophis
portoricensis (Serpentes: Colubridae). Copeia 1992, 62-68.
SAKAI, A., HONMA. M. and SAWAI. Y. (1984) Study on the toxicity of venoms extracted from Duvernoy's gland
of certain Asian colubrid snakes. Snake 16, 16-20.
SAVIDGE, J. A. (1986) The tole of disease and predation in the decline of Guam's avifauna. Unpublished Ph.D.
Thesis, University of Illinois. Champaign-Urbana.
SAVIDGE, J. A. (1987) Extinction of an island forest avifauna by an introduced snake. Ecology 68, 660-668.
SAVIDGE, J. A. (1988) Food habits of Boiga irregu/aris. an introduced predator on Guam. J. Herp. 22,275-282.
TAUB, A. M. (1966) Ophidian cephalic glands. J, M orph. 118, 529-542.
887 Brown Tree Snake Bite
TAUB, A. M. (1967) Comparative histological studies on Duvernoy's gland ofcolubrid snakes. Bull. Amer. Mus.
Nal. Hisl. 138, I-50.
VEST, D. K. (19810) Envenomation following the bite of a wandering garter snake (Thamnophis eJegans vagrans).
Clin. Tox. 18, 573-579.
VEST, D. K. (1981b) The toxic Duvernoy's secretion of the wandering garter snake, Thamnophis elegans vagrans.
Toxicon 19, 831-839.
VEST, D. K., MACKF.ssY. S. P. and KAROONG, K. V. (1991) The unique Duvernoy's secretion of the brown tree
snake (Boiga irregularis). Toxicon 29, 532-535.
WEINSTEIN, S. A., CHISZAR D., BELL, R. C. and SMITH, L. A. (1991) Lethal potency and fractionation of
Duvernoy's secretion from the brown tree snake Boiga irregularis. Toxicon 29, 4 1 ~ 7
WEINSTEIN, S. A., STILES, B. G., McCOID, M. J., SMlrn, L. A. and I<.A.Iu>oNG. K. V. (1993) Variation and lethal
potencies and acetylcholine receptor binding activity of Duvernoy's secretions from the brown tree snake,
Boiga irregularis. J. Nat. Tax. In press.
ZALlSKO, E. J. and I<.A.Iu>oNG, K. V. (1992) Histology and histochemistry of the Duvernoy's gland of the brown
tree snake, Boiga irregularis. Copeia 1992, 791-799.