= BULLETIN = = = Chicago Herpetological Society

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1 = BULLETIN = = = of the Chicago Herpetological Society Volume 29, Number 8 August 1994 The black-headed bushmaster, Lachesis muta melanocephala, is the most recently described of the four currently recognized bushmaster subspecies. In this issue Dean Ripa describes in detail captive breedings of this subspecies and of the Central American bushmaster, Lachesis muta stenophrys. Dean also provides some intriguing insights into morphological and behavioral direrences among the four bushmaster subspecies. Photograph by Dean Ripa.

2 BULLETIN OF THE CHICAGO HERPETOLOGICAL SOCIETY Volume 29, Number 8 August 1994 Reproduction of the Central American Bushmaster {Lachesis muta stenophrys) and the Black-headed Bushmaster {Lachesis muta melanocephala) for the First Time in Captivity Dean Ripa 165 HerPET-POURRI Ellin Beltz 184 Helpful Herp Hints Dennis Engler 186 NEWTLINE., Debi Hatchett 187 The Tympanum MarciaRybak 188 Photo Essay: 1994 CHS Expo Tony Rattin 189 Herpetology Unofficial Minutes of the CHS Board Meeting, July 15, Advertisements 192 News and Announcements 196 EDITORS Michael A. Dloogatch Joan F. Moore John C. Murphy Copy editor: Jill Horwich Advertising Manager: Ralph Shepstone CHS OFFICERS 1994 Marcia Rybak, President Anthony Rattin, Vice-President Gary Fogel, Treasurer Jill Horwich, Recording Secretary Brian Jones, Corresponding Secretary Michael A. Dloogatch, Publications Secretary Steve Spitzer, Membership Secretary Stacy Miller, Sergeant-at-Arms Jim Caspar, Member-at-Large Jack Schoenfelder, Member-at-Large Claus Sutor, Member-at-L^rge Ron Humbert, Immediate Past President The Chicago Herpetological Society is a nonprofit organization incorporated under the laws of the state of Illinois. Its purposes are education, conservation and the advancement of herpetology. Meetings are announced in this publication, and are normally held at 7:30 P.M., the last Wednesday of each month. Membership in the CHS includes a subscription to the monthly Bulletin. Annual dues are: Individual Membership, $22.00; Family Membership, $25.00; Sustaining Membership, $50.00; Contributing Membership, $100.00; Institutional Membership, $ Remittance must be made in U.S. fiinds. Subscribers outside the U.S. must add $12.00 for postage. Send membership dues or address changes to: Chicago Herpetological Society, Membership Secretary, 2001 N. Clark Street, Chicago, IL Publications are sent to U.S. members via third class bulk mail; the post office does not forward such mail, even if a special request is made to forward magazines. Manuscripts should be submitted in duplicate, typewritten and double spaced. Alternatively, submission of manuscripts in the form of IBM PC-compatible or Macintosh format diskettes is encouraged. Manuscripts and letters concerning editorial business should be sent to: Chicago Herpetological Society, Publications Secretary, 2001 N. Clark Street, Chicago, IL Back issues are limited but are available from the Publications Secretary for $2.50 per issue postpaid. The BULLETIN of the Chicago Herpetological Society assn ) is published monthly by the Chicago Herpetological Society. Copyright 1994.

3 Bull. Chicago Herp. Soc. 29(8): Reproduction of the Central American Bushmaster (Lachesis muta stenophrys) and the Black-headed Bushmaster (Lachesis muta melanocephala) for the First Time in Captivity Dean Ripa P.O. Box 3781 Wilmington, NC Introduction The bushmaster, Lachesis muta (Daudin, 1803), is the largest of all the vipers, and the largest New World venomous snake, reachmg a known length of 3.6 m. Rare and poorly known, it exists in undisturbed primary forests of Central and South America. Four subspecies have been described, significantly different fi-om one another in appearance. These are Lachesis muta stenophrys, L. m. melanocephala, L. m. muta and L. m. rhombeata. They range from southern Nicaragua to northern Bolivia, throughout the Amazon basin, the Guiana Shield (with the island of Trinidad), and the Atlantic forests of Brazil. The bushmaster is imique in a number of aspects. It is the only New World pit viper that lays eggs; to these it displays maternal care, guarding them protectively in its coils (no thermal incubation occurs as with the pythons, however). A long, thin, spine-shaped scale at the end of its tail produces a warning sound when vibrated against the debris of the forest floor, suggesting an ancestral relationship to the rattlesnake a snake to which it has often been compared strucmrally. It is of more slender build than the rattlesnake, however, and of a distinctly more taut, muscular appearance (older specimens, especially of L. m. stenophrys, may attain girth comparable to large rattlesnakes). It has the longest fangs of any snake in die New World (up to 39 mm in large specimens), second m length in the world only to the enormous dentition of the gaboon viper of tropical Africa. The name Lachesis muta literally "silent fate" is most apt, for bushmaster bites resuh in an 80% mortality rate in Costa Rica, even with antivenin treatment (Bolanos, 1984; Campbell and Lamar, 1989). Reproduction of two different subspecies of bushmaster occurred last year at the author's private facility. These were the black-headed bushmaster, Lachesis muta melanocephala, of southeastern Costa Rica, and the Central American bushmaster, Lachesis muta stenophrys. This marks the first time these subspecies have been reproduced in captivity (Ripa, 1994). The Dallas Zoo breeding of the South American bushmaster, Lachesis muta muta, in 1987, was the first known captive breeding of the genus. As of this writing, captive reproductions of Lachesis have yet to be repeated in any other institutional or private collections. Subject specimens here are two male and four female L. m. melanocephala collected as adults by the author in the late 1970s (when they were still classified as L. m. stenophrys) in southeastern Costa Rica; and one male and one female L. m. stenophrys collected from the wild as subadults, also by the author. The sizes of thel. m. melanocephala, respectively, are: 2.1, 2.6, 2.2, 2.4, 2.4 and 2.3 m; thel. m. stenophrys, are both 2.2 m in length. The first copulation occurred on 5 January Subsequent breedings occurred on 13 January, 7 February, 24 February (intromission interrupted), 9 March, and again as late as 23 August. The copulations were observed for their entire duration by the author, and notes taken. There is no description of copulation of the bushmaster in the scientific literature. While some examples of courtship are described in Boyer et al. (1989), the Dallas Zoo staff have not observed the actual copulations of their Lachesis muta muta (Don Boyer, pers. com., 1994). This article contains the first complete description of copulation in bushmaster^, and the elaborate and behaviorally unique rimal by which these snakes achieve it. A short explanation of the methods and materials used to achieve reproduction is included, that the reader might better understand the author's descriptions, and/or attempt similar reproductions for himself. There is a look at how these snakes show maternal care for their eggs, a subject still not well understood in this species. Greene and Santana (1983) did not observe oviposition during the course of their radiotelemetry studies of L. m. stenophrys in Costa Rica. Where are the eggs laid in the wild? Does the mother snake guard them throughout the incubation period, or abandon them after a time? This article will attempt to answer these questions. Still another curiosity about the bushmaster is its prominent spinal ridge. Here the scales attain a degree of rugosity exceeding any other species of New World snake. Does this ridge have a function in the life history of the bushmaster? An explanation will be provided for this ridge's importance to the species. Lastly, as breeding occurred of two different subspecies under identical conditions (three reproductions in all), comparisons can be made in all phases of reproduction from courtship, to copulation, to oviposition, and uhimately, hatching of the eggs. Distribution of L. m. stenophrys and L. m. melanocephala The Central American bushmaster, L. m. stenophrys, ranges from southern Nicaragua through Costa Rica and Panama, Darifen, and southerly through Pacific-coast Colombia and northwestern Ecuador. The black-headed bushmaster, L. m. melanocephala, is restricted to the remaining primary forest areas of southeastern Costa Rica, especially on, and adjacent to, the Osa Penmsula. The bushmaster's distribution in Costa Rica, as in the rest of Latin America, has shrunken markedly in recent years due to deforestation. No true picture exists as to its likely distribution today, except to say that wherever deforestation has occurred, the bushmaster has probably been eradicated. Unlike many species of snake (such as Bothrops asper), whose populations may actually increase with agriculture, the bushmaster does not seem able to adapt to the radical alterations of climate

4 and habitat imposed by the deforested environment. The bushmaster is dependent almost entirely on primary rainforest situations. Figures 1 and 2 present the known distribution of Lachesis in Costa Rica prior to 1970 (assessed from records at Instimto Clodomiro Picado, San Jose, Costa Rica), and the author's estimation of it today. L. m. melanocephala once followed Tropical Moist Forest as far west as Parrita on the Pacific coast, and as far inland as Buenos Aires, Puntarenas. It is now restricted almost totally to the Peninsula de Osa and immediate environs of the Golfo Dulce. L. m. stenophrys, once wide-ranging in Atlantic Costa Rica, has been exterminated from the entire north and northwest of that country. Its distribution once extended to within 35 km of the Pacific Coast in the extreme northwest corner. In the northeast, sporadic remaining pockets of Tropical Wet and Tropical Moist Forest, and/or Gallery Forest, may harbor isolated populations in otherwise deforested regions, but this is hard to determine. Its current distribution in Costa Rica seems limited to the Provincia de Limon, south and southeast of highway 32, where it hugs the foothills of the Talamanca mountain chain. A Brief Looli at Differences in Structure and Appearance The bushmaster's rarity, with a consequent paucity of specimens available for systematic work, has kept it from being well studied. L. m. melanocephala was not reassigned as a separate subspecies (as distinct from L. m. stenophrys) until relatively recently (Sol6rzano and Cerdas, 1986). Physical differences between these snakes, however, were long apparent to herpetologists in Costa Rica. Bolanos et al., as early as 1978, remarked differences in the immunological characteristics of the venom. Differences in appearance are rather striking. Noticeable inl. m. melanocephala are: 1.) A blue-black head dorsum, which is distinct from all other Lachesis. 2.) A bluish-grey underskin causing the scales to appear darkly outlined, especially posteriorly. This trait was not reported in the 1986 description. 3.) A very light ground color, ranging from straw yellow to cream white. 4.) Rhomboidal markings that are Wwe-black and not brown-black. 5.) Sporadic black flecking and mottled areas outside the pattern. Some strucmral differences are: 1.) Conspicuously more ventrals inl. m. melanocephala ( ) than inl. m. stenophrys ( ); 2.) Greater number of dorsal scale rows in L. m. melanocephala (37-39) than in L. m. stenophrys (34-35); 3.) Morphology of the palatine bone, which inl. m. melanocephala has a concave anterior face, and in L. m. stenophrys is flat. Interestingly, on the basis of scale counts L. m. melanocephala shows greater affinity to the South American bushmasters L. m. muta and L. m. rhombeata than to Central American L. m. stenophrys. For instance, there are ventrals in L. m. muta, and inl. m. rhombeata, roughly equal to the count in L. m. melanocephala. This similarity is hard to account for, because L. m. melanocephala is separated by at least 1500 km from the South American populations of Lachesis muta. L. m. muta is a more slender snake than L. m. stenophrys. Specimens of equal length are normally 20-30% less in mass than the latter. This is conspicuous in adult specimens, where examples of L. m. stenophrys often weigh twice as much as L. m. muta specimens of the same length. A 2 m South American bushmaster can coil within a space the size of a shoe box; a Central American bushmaster of identical age and length may require roughly twice that much space. This is not due to parasitism or diet, but appears (based on many captive-raised specimens) to be a physical difference between these subspecies. The maximum recorded length for a Central American bushmaster appears to be at least 11 feet, 7 inches (3.5 m). This example was measured by Ditmars, and was Costa Rican Figure 1. Distribution of Lachesis muta stenophrys (Atlantic coast) and L. m. melanocephala (Pacific coast) in Costa Rica prior to Figure 2. Current distribution of Lachesis muta in Costa Rica since deforestation. 166

5 in origin. There is reason to believe that South American bushmasters may exceed this length, reaching a length of at least 4 m or even more (Theo Henzen, Paramaribo, Surinam, pers. com.). Hoge and Lancini (1962) suggest thati. m. muta may reach 4.5 meters. However, given the weight differential between these two subspecies, the South American form might not appear to be as large when coiled. A detailed look at the Lachesis muta subspecies shows numerous other differences morphologically particularly in the size, shape and scutelation of the head. Relative to body length, the head of the Central American subspecies is considerably larger than the head of its South American counterpart. In an L. m. melanocephala of 120 cm (a yearling), the head is 58 mm from the angle of the jaw to the ventral, 40 mm across laterally, and 29 mm diick at its greatest vertical point. This is approximately equal to the head size of a two-year-old L. m. muta of 30 cm greater body length. The heads of South American bushmasters may run as much as 15-20% smaller in proportion to body length than Central American ones. This ratio to head size by no means follows only with juveniles; rather it seems consistent throughout the development of these snakes (see Figure 3). To the audior's knowledge, this important relative difference between the subspecies has not been recorded in the systematic literature. A relatively different head shape is another facet that has mysteriously escaped the eye of taxonomists. A look at Figure 3 will show how appreciably different head shape can be between Central and South American specimens. In the dorsal view (Figures 3e and 3f), the head of L. m. stenophrys (and L. m. melanocephala, not depicted) appears oval, and blunt at the snout, with the supraoculars set relatively far forward. Compare this to the dorsal view of L. m. rhombeata (nearly identical in shape to L. m. muta): The head is distinctly more spadelike, relatively pointed, and the snout somewhat elongate. At a glance one could mistake it for the head of a Bothrops species. In the lateral view ofl. m. stenophrys (Figure 3g), the head is thick and boxlike, with a squared off snout, rostral nearly flush with mental. The eye is small, set relatively far forward on the snout. The loreal pit seems relatively close to the eye. The lower jaw is prominent, and appears acutely tilted forward anteriorly. If the head of the South American form resembles a Bothrops, the Central American form gives the impression of a trend toward Crotalus. In the lateral view of the South American form (Figure 3g), the eye appears larger, more protuberant in the relatively smaller head. The eye is relatively farther behind the loreal pit and the snout. The snout appears longer. The whole head seems flatter, more compressed vertically. The chin, at the mental, appears relatively recessed beneath die rostral, with the infracephalic region less prominent beneath the supralabials. The entire lower jaw appears more compactly situated in the head. To what extent are these differences reflected osteologically? The author can find no data. Such phenomena seem not to have been addressed in the systematic literature. Morphological scale differences will be apparent even to the untrained eye. To mention a few: The prenasals of L. m. muta and L. m. rhombeata may be enlarged, protuberant, triangular and angled anteriorly; but inl. m. stenophrys and L. m. melanocephala these are much reduced, with a flat or rounded face. The internasals in L. m. muta and L. m. rhombeata may be enlarged, in contact, or separated by one or two small scales; but in L. m. stenophrys and L. m. melanocephala the internasals are much reduced, in some specimens so small as to be undetectable among the numerous small scales of the prefrontals. The canthals of L. m. muta and L. m. rhombeata are elongate, distinct and upraised; in L. m. stenophrys and L. m. melanocephala, they may be more oval, indistinct, subdivided, flattened or smooth. The canthus rostralis appears relatively broader and more rounded in the South American subspecies, where the higher canthals form a small ridge lacking in Central American specimens. Of immediate evidence in South American forms is the opening to the nostrils, which, because of the more exaggerated prenasal, and the concave, and often divided postnasal, appear more conspicuous in the two South American subspecies (especially in L. m. rhombeata). By contrast, in the Central American subspecies, the actual openings to the nostrils appear extremely small, to the degree that in some specimens the nostrils may be hard to detect. There are a consistently higher number of supralabials in South American examples than in diose of Central America: 8-11 in L. m. muta and L. m. rhombeata, and 7-9 in L. m. stenophrys andl. m. melanocephala. In Central American examples the supralabials are usually strongly crenelated at the border of the mouth; in South American examples they may be relatively smooth along this border. Interoculabials and lacunolabials may be highly variable from Central American to South American examples as well. Campbell and Lamar (1989) state that "the dorsals [of Lachesis muta] are broad with rounded apices and lack apical pits.... The free apical portions of the dorsals are meager...." The confusion generated by the subspecific nomenclature may be illustrated: Campbell's and Lamar's reference evidently emanates from the prevailing monotypic classification with its assumptions of certain uniformities existing between these snakes. For, by contrast, in some adult bushmasters the dorsal scales can be more elongate than broad, with free apical portions moderately developed, and with well defined keel pits. The author remarks an atypical L. m. melanocephala that exhibited all these characteristics in one specimen. Keel pits are sometimes seen in L. m. rhombeata, and in older, highly keeled examples of L. m. muta. Again, from Campbell and Lamar (1989), "... the height of the keels diminishes laterally and the first 3-5 scale rows may be smooth." This is certainly true of adult L. m. stenophrys, where this smoothness may be displayed even to the sixth scale row and above; but in some adult specimens of L. m. muta, L. m. melanocephala, and even relatively young L. m. rhombeata, the keeling may diminish much less laterally, remaining strongly keeled even to the second or first scale row. The beaded, tubular keeling of die middorsals in L. m. muta, and L. m. rhombeata far exceeds in height that of L. m. stenophrys, and to a degree, L. m. melanocephala. A look at 167

6 a) Head of L. m. muta, showing greater elongation and vertical compression of South American Lachesis forms. Tends towards a Bothrops model. b) Head of yearling L. m. melanocephala, showing the thick, blunt head of Central American forms. Tends toward a Crotalus model. Note simplified scutelation, conspicuously different from L. m. muta in adjoining photo (see text for details). c) This two-year-old L. m. muta has a more rounded, less laterally compressed body cylinder than is found in the Central American forms. The head is relatively pointed, like a Bothrops. Note large, broad, evenly distributed dorsal scales; free apical portions are meager. d) L. m. melanocephala yearling, showing strong lateral body compression. The snout is blunt, the whole impression rattlesnakelike. The dorsal scales are smaller, more elongate, less evenly distributed, with free apical portions more conspicuous a trait that will be retained in adulfliood. This yearling has not yet undergone ontogenetic color change. The paravertebral stripes and broken vertical barring are not uncommon in Central American specimens of both subspecies. e) Dorsal head comparison of bushmasters from opposite ends of their range: Central American L. m. stenophrys (left), and South American L. m. rhombeata (right). Head of Central American bushmaster is large, oval and blunt. Head of South American example is lanceheadlike, pointed. Note head size difference; these females are of nearly identical TL: 1700 mm and 1630 mm respectively. Arabesque-like head markings are characteristic of L. m. rhombeata. i ' i i 11 J i J 1 1» i I f) Detail of dorsal 3 head 14 view. 15 Note the 16 internasals ir and 18 prenasals: 19 enlarged in L. m. rhombeata (right), and reduced in L. m. stenophrys (left). These trends are shared respectively by South American L. m. muta and Central American L. m. melanocephala. See text for some other scale comparisons. Figure 3. Four subspecies? Or three distinct species of a single genus, one with two subspecies? The author suspects that the longstanding accepted classification of the bushmaster may be a little too conservative. Here is a photographic look at four forms of Lachesis muta. All photographs by the author.

7 g) Lateral view of same examples of L. m. stenophrys (upper) and L. m. rhombeata (lower). Head of Central American form is distinctively blunt, oval, with small eye set above large supralabials. In the South American specimen, note relatively larger and more protuberant eye with its more vertically compressed head and more pointed snout. h) Ventral view of same L. m. rhombeata (upper) and L. m. stenophrys (lower). Numerous differences in infracephalic scales. Oval Central American head form is retained ventrally, while the South American form retains significanuy greater resemblance to lancehead snakes. i) Ventral view of heads of adult female L. m. muta (left), 2131 mm TL, and L. m. stenophrys (right), 1720 mm TL. The L. m. muta specimen is a full 410 mm longer than the L. m. stenophrys and still the head is relatively smaller up to 20% smaller by length. Lancehead-like morphology of South American head form is retained. (The dorsum of diis L. m. muta specimen is regrettably disfigured, and has not been included). j) Lateral view of markings of the four subspecies. Soudi American forms, upper (L. m. muta and L. m. rhombeata), and Central American forms lower (L. m. stenophrys and L. m. melanocephala). Blotches of South American subspecies retain rhomboid shape laterally. Central American forms tend toward vertical barring. Note stronger lateral body compression in Central American specimens. k) Dorsal view DI markings ot ihc lour suli^pecies (same order as previous photo). South American forms show significantly less lateral body compression. 1) Dorsal view of live Central and South American examples. L. m. muta male (smaller), and L. m. stenophrys gravid female (larger). Examples are 1900 mm (L. m. stenophrys) and 1750 mm (L. m. muta) of nearly equal length. Note conspicuous difference in girth, head size and head shape of the relatively more slender L. m. muta. Central American bushmasters (all apart from this specimen's gravidity) are consistently more robust than South American ones. Figures, (cont'd) 169

8 Figure 3 will show how different the shape of the dorsal scales can be between Central and South American forms. Note the even, almost geometric dorsal scale distribution of the broadscaled Surinam L. m. muta, as compared to the more irregularly distributed, smaller and elongate dorsal scales of the L. m. melanocephala pictured (Figures 3j and 3k). The osteology of Central as opposed to South American forms reveals other trends, not the least important of which is the relative height of the vertebral ridge. This ridge (so unportant in the life history of the bushmaster, as we shall see) varies considerably between specimens from Central and South America. In both Central American subspecies the ridge is highly developed, but in Surinam L. m. muta the height of the neural spines may be much lower, so that the ridge appears only marginally developed, and in some specimens not at all apparent except anteriorly (Figures 3c and 3d). This variation is curious and inexplicable. In examples from the lower Amazon Basin, and from Mato Grosso, state of Brazil, the vertebral ridge is once more in very high relief, comparable to examples from Central America. L. m. rhombeata consistently exhibits a pronounced middorsal ridge throughout its distribution range. Perhaps of greater significance is the relative height of the body cylinder in Central American as opposed to South American forms. Throughout its distribution range, L. m. muta shows less lateral body compression than Central American forms, to the degree that the midthoracic body segment appears relatively more round. This seems not entirely due to the less prominent ridge of some South American specimens. Certainly this could be a promising area for exploration as regards the evolutionary divergences of these large crotalids. The above descriptions reveal a geographic pattern to the systematic evaluation of Lachesis. A more incisive comparison of Central and South American forms could doubtless yield many more differences that have not been accounted for by taxonomists, but which, perhaps owing to the relative difficulty of havmg sufficient examples of all four subspecies for smdy at any one place and time, have escaped systematic remark altogether. The subspecific classification of the bushmaster, with one exception, occurred during the middle part of this century, with examiners geographically isolated from each other by many thousands of miles across the largest part of South America and part of Central America (most work was done at Instituto Butantan in south Brazil, and Instituto Clodomiro Picado in Costa Rica). In those days the finding of any bushmaster was a rare and extremely noteworthy event. Well-preserved specimens of Lachesis were highly prized and not easily released to other institutions. Exammers often had to rely on written descriptions from other laboratories to do comparative work. To complicate things, postal correspondence between points in South America required many months. As a result, the determination of subspecies in the bushmaster was made ahnost entirely on the basis of what could be easily written down, that is scale counts. Beyond these, and notations as to color and pattern, physical comparisons between bushmaster subspecies (lightly touched upon here), were evidently never made at least they have not come down to us in the literature. The modern examiner, wishing to attempt such a re-evaluation for himself, should bear in mind that ontogenetic scale changes in the bushmaster can be severe and misleading, and that only examples well into adulthood provide a consistent view. Dorsal body patterning of bushmasters, formed of dark diamond-shaped blotches, rhombs, or saddles, is also relatively different in Central American as opposed to South American subspecies (Figures 3j and 3k). Regional color differences between bushmasters may be more radical than was previously believed. The ground color of L. m. muta can be a light ciimamon orange or coral pink, but in other examples it can be as uniformly brown as that of a cottonmouth moccasin {Agkistrodon piscivorus). Conversely, L. m. sterwphrys, usually described as a darkly colored race, exhibits a fair number of specimens with light and strongly contrasting ground color (see photo comparison to L. m. muta. Figure 31). L. m. melanocephala reaches a very light straw yellow in adulthood; at least five adult examples in the author's collection are uniformly cream white offsetting the blue-black dorsal markings. Mr. Earl Turner, who has had experience with many bushmasters over the years (and has survived ttie bite of a 2.5 ml. m. stenophrys), reports specimens of L. m. muta from near Cochabamba, Bolivia, as having a lemon yellow ground color comparable to yellow Bothriechis schlegeli (Earl Turner, pers. com., 1994). Bushmasters of all subspecies tend to be uniformly dark in color when young (up to their first and second year), evidently a survival advantage. Central American bushmasters of both subspecies appear to gain substantially more black pigment in the dorsal markings after their second year, with the ground color becoming conspicuously lighter and more contrasting. These ontogenetic color changes are usually complete by the time the snake reaches sexual maturity. Nomenclature The above section serves to illustrate the foibles of the typical view that a bushmaster is basically a bushmaster regardless of where it comes from. The author has cited weight differences, body elongation, head size, head shape, scale differences of the head and body, even the general shape of the body cylinder. With greater examination the subspecific nomenclature begins to look more and more inadequate to describe these taxa. In the light of recent taxonomic assignments afforded to systematically less distinctive forms (e.g., Agkistrodon piscivorus subspecies), the current nomenclature afforded to Lachesis muta presents an incongruity if not an absurdity. When determinations between two full species, for example, Bothrops leucurus and B. pradoi, known to intergrade and predictably interbreed can only be resolved by an expert with a microscope, snakes so obviously divergent as the Lachesis muta subspecies which have no intergrades begin to seem gigantically mislabeled. Having advanced some systematics, the author wishes to interject a strong statement based on a more holistic view than that of the assemblage of details: No eye practiced at viewing the forms of snakes, once having had an opportunity to view all these animals alive and 170

9 together at one time, can ever come away with the impression of having seen a single subdivided species. There are other inconsistencies difficult to resolve for the examiner. A thorn in the side of current Lachesis nomenclature (one to my knowledge, never resolved) is the curious case reported by Martinez and Bolanos (1981), of a South American L. m. muta (or what appeared to be one) found in the Darien region of Panama, Central America. The Isthmus of Panama falls within the range of L. m. stenophrys, not L. m. muta. The equivalent, in terms of geologic barriers, of finding a L. m. muta less than 200 km from the Panama Canal, would be to find a western diamondback rattlesnake {Crotalus atrox) in Florida. The specimen, which was deposited in the Zoological Museum of the School of Biology of the University of Costa Rica (No. 8061), matches written descriptions of L. m. muta entirely and yet, is it? Campbell and Lamar (1989), who cite the report as a source, remark of such specimens, "... we suspect on geographic grounds that their affinities are with L. m. sterwphrys." If this is so, then the question has not been settled, only rephrased: What variafions of L. m. stenophrys exist that specimens so nearly approximate to L. m. muta can be found in Panama? The weakness of the current nomenclature is thus evident it does not permit acknowledgment of regional structural variations within the existing subspecies, and hinders description. This uhimately creates a blank spot in these areas of study. The fact that the Martinez specimen exists at all suggests that subspecification may be possible widiin the "subspecies." The current nomenclature offers no means to resolve such problems. The taxonomy of the nominate Lachesis muta muta is much more dramatically afflicted. L. m. muta's distribution range is enormous, in fact, one of the largest distributions of any snake species in the world over 1,500,000 square miles. Within this hugely varied range, L. m. muta presents an amazing homogeneity of forms; yet sufficient morphological differences exist to divide, and perhaps subclassify, many regional populations; e.g., those of the Guiana Shield, and those of Mato Grosso, Brazil (to name two differing types which the author has examined). The nearly analogous (to Mato Grosso specimens), Lachesis muta rhombeata, holding to its original assignation by Hoge, is accurately not more than a subspecies or variation on what is now called Lachesis muta muta. It has the same flat, elongate head, relatively larger eyes, and shares the prominent prenasals and internasals of L. m. muta. It shares nearly identical patterning to some Mato Grosso L. m. muta (except being of more brilliant hue), even to the elaborate arabesquelike head speckling. A more descriptive way to classify the bushmaster might be to acknowledge the two Central American forms as distinct species of the genus Lachesis, distinct both from each other and the nominate South American form. This is not only a more accurate assessment of their physical differences, but it will allow for taxonomic distinctions not currently possible within at least two of these groups {L. m. stenophrys and L. m. muta). This may ultimately lead to a more complete understanding of the origins of these snakes, their morphogenetic ancestors and descendants. Lachesis, because of its egg laying, has always presented smmbling blocks to phylogenetic theories. It seems evident that at some point the Central American forms began to evolve in a different direction from those of South America. Why? To what species did they tend? At some point a common ancestry must have occurred; what were their common, or perhaps uncommon descendants? The author has collected examples of all four of the currently assigned subspecies in the wild {L. m. stenophrys, L. m. melanocephala, L. m. muta andl. m. rhombeata), and at present possesses live specimens of each. His conclusions here are based on these live references, and on preserved ones both here and at Instituto Butantan, Sao Paulo, Brazil. There has been very little in the way of comparative data written about these snakes. At present, the comparative taxonomy of Lachesis consists largely of a handful of relatively disparate looking drawings from countries widely removed geographically, with examiners all quoting each other but only the most general data base to go back to for comparisons. A dearth of available specimens for examination by taxonomists will likely hold up any revision for many years to come. To avoid confiision, however, in this article all further references to bushmaster subspecies will be made using the current nomenclature. Function of the Black Head Dorsum in L. m. melanocephala, and Its Relation to High Elevations In Costa Rica, the southern central mountain range (Talamanca) constitutes a strong geographic barrier that separates Atlantic and Pacific populations of Lachesis muta and other snakes. This barrier, reaching altitudes of m, prevents intergradation between L. m. stenophrys and L. m. melanocephala, so far as is known (Sol6rzano and Cerdas, 1986). L. m. melanocephala has been collected from substantially higher elevations dian have been recorded for L. m. stenophrys in Central America as high as 1500 m in Premontane Wet Forest-Rain Forest transition (Solorzano and Cerdas, 1986), some 500 m higher than has ever been noted for L. m. stenophrys. This is equal to the highest records that exist for Lachesis m. muta in South America, where nearness to the equator makes for somewhat greater warmth than in Central America. Temperamres at elevations of 1000 m can average between 12 and 20''C year-round a "frigid" climate for large ectotherms, and especially egg-layers like the bushmaster (Vial etal., 1967). Observations by the author suggest that L. m. melanocephala may be more diurnal than other bushmasters. Here, at this facility, L. m. melanocephala regularly emerge from their simulated burrows in midafternoon (about 1400 h EST). They have the extraordinary habit never before reported for the bushmaster of periscoping the head up from the burrow, and holding it elevated and fixed in order to bask their black head dorsums in sunlight commg through a window. This is achieved while keeping the rest of the body submerged and concealed within the burrow, extended to its full body length (for a description of the simulated burrows by which these observations were made, see "Materials and Methods of Husbandry and Breeding"). In this position, head held in midair 171

10 some cm above the ground, the snake may remain nearly motionless for up to two hours. On Isla Violma in Costa Rica in 1986, the author observed a 2 m male L. m. melanocephala holding its head elevated in sunlight for over 30 minutes, while keeping its main body completely hidden in a paca (Agouti paca) burrow. As darkness fell, the snake emerged entirely. Thus the purpose of die black head seems to be thermoregulatory. Relationships of body size and conditions of meeting thermal requirements among ectotherms are well known. In the higher elevations inhabited by this subspecies a period of head basking before dark could be important for this large ectotherm to remain active during the cooler temperamres occurring after nightfall. By protruding only the head in this way, the snake does not expose itself to predators. By contrast, of 12 adult L. m. stenophrys maintained at this facility formerly, all were strictly nocturnal, emerging usually after about 2000 h. The author has never observed L. m. stenophrys to bask in sunlight, neither their heads nor any parts of their bodies. Nor were any of the three male and seven female L. m. muta that were maintained at the author's facility in the mid-1980s ever observed to bask in direct sunlight. Dallas Zoo, however, reports some basking of gravid L. m. muta (Boyer et al., 1989). It may be that basking in bushmasters (other than L. m. melanocephala) is normal only to gravid snakes. Aggressive Behavior of L. m. melanocephala The bushmaster's reputation for aggression against man is of almost mythological proportions. Accounts of its attacking and actually chasing intruders are at variance with the reports of many herpetologists, however, who remark more usually of its placidity, even timidity. "Deceptively lethargic," and "calm disposition," have been remarked of L. m. muta andl. m. stenophrys in captivity (Burchfield, 1975; Boyer et al., 1989). The seeming sluggishness of some bushmasters when encountered during the daytime may present a misleading picture. And bushmasters often "tame" readily in captivity. Equally, mishandling (i.e., "noosing" or other forms of bodily restraint), seems to put specimens in a permanently benumbed state. This is typical of imported specimens that have been shocked or injured by capture. "If restrained, it fights desperately and once captured it will sulk in a comer of the cage for days, never moving from the spot, not even to approach a dish to drink." (Ditmars, 1937). Under such conditions, or when deprived of significant sensory cues, such as those chemical markers placed by the snake to orient itself within its own environment (such deprivation occurs when removing the animal to a novel environment, or through complete removal of a disposable substrate like newspaper), this normally active anunal enters a state of lethargy and torpor; it appears to have trouble crawling, coiling or even holding its head up. In its own enviroiunent, however, or when kept under optimum conditions, the bushmaster will usually defend itself readily. Threat displays may include audible tail vibration, inflation and stiffening of the body, and inflation of the throat/gular region (especially L. m. muta). Defense reactions may involve striking from an elevated head stance, while the body shifts itself in a fluid, moving "S" coil toward the intruder. L. m. melanocephala has a more nervous, irascible disposition than other bushmasters. It is capable of aggressive behavior that at times may include chasing the intruder from its immediate vicinity. At San Pedrillo, Costa Rica, near Corcovado Park, an adult male L. m. melanocephala pursued the author a distance of some 8-10 m and left a safe burrow to do so. Whereas L. m. stenophrys andl. m. muta normally stand their ground and strike from a fixed coil, L. m. melanocephala will sometimes advance bodily toward the keeper, its head upraised to a level of 90 cm or even more above the ground. Such behavior, however, seems always associated with the breeding season, when specimens become particularly active and bold (examples will be described). Is the aggressiveness of L. m. melanocephala tied to its being more diurnal than other bushmasters (owing to its head basking), and thereby necessitating a greater ability to defend itself against predators? Behavioral differences The author has observed numerous behavioral differences between these subspecies. The following brief notes, expressed in relative terms, will show some of these differences, as well as some of the basic traits common to the genus. The author recognizes that all these may be subject to exceptions and conditions under which observations took place. L. m. melanocephala commonly crawls with its head strongly elevated, cobra-like, to a height of 50 cm or even more above die ground; nocmrnal (in feeding), but partially diurnal (see "Function of the Back Head Dorsum..."); active, nervous, bold when confronted, some examples show little fear and will approach the intruder (this need not be taken for signs of aggression, however); in defense, may elevate head from a fixed coil like Crotalus durissus, or advance toward the intruder by a series of shifting "S" loops; some examples may press home to attack (usually females), or bluff a lunge and then rapidly mrn and flee (males); ambush hunts on open terrain from a fixed coil with elevated head, but will also actively pursue prey for short distances (for a brief description of ambush hunting of L. m. stenophrys, see Greene, 1983); a good and frequent climber to considerable heights (to above 3 m); strike-holds prey ahnost exclusively, only rarely pursuing dying prey after strike-release. L. m. stenophrys is less prone to elevate the head while crawling (seldom held above 30 cm); very nocmrnal; normally calm, but males nervous and defensive during breeding season; a hard, heavy striker, may strike hard enough to thrust its whole body forward, and need not necessarily coil to do this; this strike may come unexpectedly, even before a warning "buzz" from the tail has been sounded; defends itself from a fixed coil and may remained thus positioned for hours if the intruder remains present; never observed to advance to attack, but seldom flees eidier, at most usually turning and crawling slowly away, vibrating the tail, while keeping the head directed toward the intruder; an infrequent climber though capable, usually avoids climbing to any real height; ambush hunts with 172

11 head slightly elevated above coils (seldom to the elevation seen in L. m. melanocephala); will sometimes pursue prey to an area within striking range, take a fixed coil and wait for prey movement; prefers mammal prey almost entirely, but will take birds; normally strike-holds prey but will strike-release and trail envenomated prey for short distances. L. m. muta seldom elevates head to any considerable degree while crawling, rather it seems to do just the opposite, feeling its way along with head held low as if in an attempt at concealment; crepuscular, strongly nocturnal, in the wild most active after midnight; generally more cryptic than Central American subspecies light seems to have somediing of the effect on them that rt has on a Bungarus the sudden appearance of light causes them to "freeze" in motion; as a resuh, in daylight, some specimens give the impression of docility in that they can be approached quite closely and even picked up bodily without employing defense postures; unpredictable in defense, may be shy, hiding head under the coil, or may bolt quickly and flee; young specimens typically defend themselves vigorously; often gapes mouth in defense posmre (less often seen in other subspecies); whereas L. m. stenophrys may take an elevated head stance, rearing the neck back into order to launch a strike, this snake seems to favor concealing the head, pressing the nose against the ground, and then launching the strike out from behind a slightly upraised coil; sometimes bluff strikes with mouth closed, or open mouthed with fangs not engaged, but meaningftil bites occur; capable climber into thickets; fond of birds as well as mammals; will actively follow prey for short distances; does not hold head very elevated when ambush hunting; strike-holds small prey, but strikereleases prey beyond a certain size threshold, and this Uireshold is generally much lower than for Central American subspecies; will pursue dying prey for short distances. On L. m. rhombeata I have accumulated very few observations as of this time. They are seemingly much more prone to a meaningftil strike-bite than L. m. muta; will lunge at or alternatively flee the intruder; strike-holds and strike-releases prey; seems very similar in many respects to L. m. muta. All of the above exhibit typical bushmaster defense posnires to varying degrees (see "Aggressive Behavior of L. m. melanocephala'" for a description of these). Materials and Methods of Husbandry and Breeding 1) Because the bushmaster is a large snake, and very active during courtship, considerable space is required for the ritualized movements of the male and female. The enclosures used here were approximately 4.2 x 3.6 X 2.4 m high, forz.. m. melanocephala; and 3.6 x 1 x 1 m high, forl. m. stenophrys. Regarding the much narrower size of the second enclosure, in which breeding occurred successfully also, length seems to be more important than breadth, allowing the snakes to stretch out to their full length during courtship. Any enclosure in which the keeping or breeding of Lachesis is attempted should have at least a reasonable height also (1 m or better); this is very important for L. m. melanocephala, for this subspecies commonly crawls with its head upraised to a height of cm or more above the ground floor. 2) The enclosures have a substrate of cypress mulch filled to a depth of about cm. The mulch, when moistened, maintains humidity. The larger enclosure is equipped with artificial rain systems which can be controlled from outside. The smaller enclosure is not so elaborately equipped, and must be irrigated with a hand-held sprayer. Hiding places in the large chamber consist of six simulated underground burrows of 2 m long X 45 cm wide corrugated pipe. These tunnels, designed to imitate paca burrows, allow the snake to extend itself to near full body length, and yet remain unexposed. The tunnels are partially submerged in the mulch substrate, and filled to a level of about 10 cm with peat moss (the moss retains humidity). Artificial rain systems capable of misting (or flooding) the burrows allows adjustment of humidity levels within the pipe. Sexual activity of bushmasters has been observed here to have been associated with raining on their skin. In the smaller chamber, one 1.6 m long corrugated pipe 45 cm wide is employed as a hide area for a single pair of snakes, this situated outside the cage in such a way that the mouth opens into it; this prevents much needed level and open space being taken up in the enclosure. This tunnel contains no misting or raining apparatus. Reproduction occurred equally well in both chambers, substituting the hand-held sprayer in the smaller one bearing in mind diat L. m. stenophrys is characteristically a much less temperamental anunal flian L. m. melanocephala. The latter often cannot tolerate much stress of keeper contact. Also, the L. m. stenophrys specimens were collected as juveniles, more readily adaptable to captivity than adults. 3) The normal ambient temperamre for these enclosures is C (see Table 1). A gradual increase of ambient temperature prior to the breeding season, to 25 C, with the withholding of humidity (to below 50%), was intended to simulate a shortened dry season in Costa Rica (November-December). This temperature increase and reduced humidity was maintained for one month (20 November through 20 December). Starting in late December, the rain systems were used, creating a humidity increase to above 95%, with the temperature increase maintained. This is quite different than the method employed at the Dallas Zoo (see Boyer et al., 1989). At Dallas Zoo, through February, March and April, temperamres are lowered to 23.9 C for several days, and returned to C described as the normal temperature of dieir enclosures (this is 5-7 degrees higher than the temperature normally employed by the author). In the author's enclosures, a dry season Table 1. Temperature and humidity regimes to maintain and breed bushmasters in captivity. Normal year-round photoperiod = 12 h/12 h. Temperature Relative humidity (%) ( C) Normal ambient November - December January - March 0000 h h 1230 h h

12 is simulated, temperature is elevated, and then a mock "rainy season" synthesized with appropriate humidity increase and raining on the specimens. Temperature is allowed to vary during die day, elevated in the afternoon and early evening, then allowed to fall after midnight, to the normal ambient C. The photoperiod is 12/12 h year-round, coinciding to the near equatorial photoperiods in areas inhabited by these subspecies. COURTSHIP First Signs of Sexual Activity Signs of reproductive interest in diese snakes were first observed in female L. m. melanocephala 3, who switched her tail about when sprayed with water. This specimen had been observed coiling outside her burrow more often than usual, for a period of about 40 days (November-December 1992). In mid-december male L. m. melanocephala 1 showed a high tongue flicking rate (TFR) in the area of substrate where female 3 had coiled. This TFR was estimated to be above 50 flicks per minute. He crawled over this area repeatedly, coiling up there for short periods of time. Within the next three days this male frequently explored the enclosure. He ceased his regular pattern of hunting activities. No ftirdier interaction with this particular female was ever observed, however. In late December another female, L. m. melanocephala 2, was observed slowly crawling die perimeter of the enclosure. She dragged her posterior body over die substrate, tail upraised vertically, vent partially open, contracting her cloaca at frequent intervals. This activity was almost exclusively diurnal. It began at 1600 h for 2-3 days, and then gradually earlier, at about h for about seven more days. The presence of full daylight in the room did not seem to impede her. (It is very unusual behavior for these snakes to completely emerge from the burrow earlier flian 1500 h; die diurnal Figure 4. L. m. melanocephala female awaiting rendezvous with male. Temperamre/humidity regulator indicates ideal breeding conditions. Note strongly oval head shape of Central American forms. Scale distribution and morphology are unlike those of L. m. muta, tending more toward those of allopatric Central American L. m. stenophrys. This specimen is uniformly creme yellow with blue-black dorsal markings. Photograph by the author. head basking of L. m. melanocephala has been described). The female continued this process, crawling back and forth over about a 5 m^ area, each afternoon for a period of about 3 h. Food offerings (as live prey) were refused by this specimen during this period. By 1800 h she had retired to her burrow. No breeding, however, occurred at this time. On 1 January 1993, after a period of several days, yet another female, L. m. melanocephala 4, began showing signs of similar activity. At about 1640 h, she began frequent crawling about the enclosure. Upon encountering the crawling pattern of female 2, she displayed virtually identical TFR to the previously described male's. She touched her chin repetitively against the substrate as she crawled, with rhythmic bobbing movements of her head and anterior body. (These movements were similar to those that would later be observed in these specimens during courtship.) As female 2 was also active during this time, die two female snakes eventually met up, and began crawling against die sidewalls of the large cage together. They "chinned" each other's dorsums widi repetitive, bobbing head movements, and momentarily intertwined their bodies. Their tails were upraised vertically as they crawled. Cloacal contractions were observed. The keeper inferred that they were chemically scenting die substrate. A particular area of the enclosure was then focused upon for their activities. They crawled the same 5 m^ area of substrate with tails upraised, five to six times. After a period of about 2 h, TFR was reduced, and finally subsided. The two snakes separated and returned to their individual hiding tunnels. This activity of the two females was repeated each afternoon until 4 January, three days later. On 4 January, at about 2000 h, L. m. melanocephala male 1 emerged from his hiding tunnel to excrete feces. Upon encountering die area of substrate frequented by the females, his TFR increased markedly (from 20 to 60 per minute). Widi his rostral pressed lightly to the cage floor, he made repetitive bobbing movements of his head as if tracing an unseen but specific pattern left on the substrate. This was followed by a period of repeated crawling over that area. At times he would coil, wait for some minutes, neck and head portion extended to a length of 30 cm and held suspended some 20 cm over die substrate. Then he would crawl out of his coil and repeat his circuit of die immediate area. Rapid TFR with the tongue touching the ground surface was observed. The next day the author noted a whitish secretion, consisting of two milky globules spaced approximately 30 mm apart on die substrate, coinciding with what would be the position of the male snake's hemipenes. This was deduced to be a sperm plug. It was not removed, but allowed to decay on the mulch substrate. Neither of the two females appeared from the burrow on diis day. Food was refused by all female specimens during this period. In males, feeding occurred less regularly, but was not discontinued. 174

13 First Copulation of a L. m. melanocephala Pair On 5 January 1993, at 1600 h, courtship began between male L. m. melanocephala 1 and female L. m. melanocephala 2. Temperamre was 24 C, elevated from 20 C diat morning, with a humidity above 95%. This was one of the most violent couplings the author has ever witnessed in snakes. It included the male acmally knocking the female off the ground into the air with his coils. An impressive display among snakes that exceed 2.2 m and 6 kg. Thrashing and violent sounding movements continued in the cage throughout the evening, well audible from die next room. At times the female would form a solid coil, withdrawing her tail and lying on top of it; a position which offered the male no access to her. At intervals she would hide her head under her coils. The male would advance rapidly to her, chinning her dorsum, and then and this was most extraordinary behavior for a snake he wouldover upside down on his back on top of her, and coiling there, venter upwards, commence a rapid backward/forward sawing of his dorsal body against her (Figure 5). In this incongruous belly-up position, the male maintained the rapid fiddling of his coils for some 60 seconds. It was as though he had converted his whole body for a few moments into a flexible "zigzagging" saw. The rasping noise of his sharp dorsal scales against the female's dorsum were audible from a distance of some 10 m. The laterally compressed body cavity of the bushmaster, resulting in a pronounced dorsal ridge of exceedingly hard and sharp lepidosis, has long been a cause of wonder to herpetologists. No evidence for what the bushmaster might use this ridge for has been given in literature, aldiough some function for it would follow. Only the Neotropical rattlesnake {Crotalus durissus) displays a comparable spinal ridge, or comparably rugose scutellation, in New World snakes but the condition is even more developed in the bushmaster. Here die middorsal scales are prominent, highly mbular, and of almost knifelike character, to the touch feeling ratfier like the edged teeth on a wood rasp. In older adult specimens these scales can resemble small tubular spikes, sharp enough to inflict skin lacerations on anyone unwise enough to attempt restraining one with bare hands and arms (this happened to the author in 1985 while hand collecting a 2.2 m adult L. m. muta). In Surinam (Tepoe, District of Sipaliwini) in June 1986 die author observed an L. m. muta using this ridge in a "sawing" motion to sculpt and excavate the overhead portion of a passage in a partially demolished paca burrow {Agouti paca), so as to create space to coil (Ripa, 1994). But here the design was revealed to have another function as well a* a means of sexually stimulating the female. Such interesting behavior as "upside-down sawing" during courtship has never before been reported for any snake species. It is evidently a behavior unique to the bushmaster. It is typical of all the breedings that took place at the author's facility, common to males of both L. m. melanocephala and L. m. stenophrys, and was observed on six different breeding occasions from January to mid-august Whether this characteristic is shared by South American L. m. muta and L. m. rhombeata is not known. Failing to break apart the female's coil, the male righted himself. Placing a bent coil against the female's flanks, he suddenly straightened himself The result was like a punch that knocked her bodily off the ground. The female fled at once, throat ventrally inflated, and tail vibrating loudly (note these are defense responses). The male quickly pursued. Again finding her coiled, he repeated the maneuver of upsidedown sawing on top of her. Righting himself, he struck her widi the outermost side of his coil once more. Again she was knocked clear from the substrate. At last, loosely coiled, die female was manipulated to the full lengdi of her body. Sprawling along her flanks, the male attempted to wind himself about her, while continuing the bobbing motions of his chin and anterior portion against her nape. By 1830 h their tails had wrapped, and vents were adjacenfly positioned. Here the male used his tail as though prehensile, and bent the female's tail back at an angle toward her main body. The effect was to part the female's vent. During this period, her posterior body, for about 10 cm anterior to the cloaca, became visibly engorged. By 1855 h her cloaca had become widely dilated, opening to a diameter of approximately 30 mm. A reddishbrown secretion emerged and dripped onto the substrate. It did not appear to be of fecal material. Intromission was accomplished by 1935 h. The hemipenis was plainly visible throughout much of the process, partially exposed from the vent of the female. It was of grayish-pink coloration, and approximately 15 mm in diameter. Calyces were visible at the lobes, and rings of spines. A secretion was left on the substrate. Throughout die copulation, die male continued the vibratory motions of his head and neck upon the female. Rapid TFR (above 50 per minute), and twitching and chinning of the male's anterior body, continued at intervals until approximately 2330 h. Eventually, the female's anterior portion became turned opposite to the male's, so that they faced in different directions. When this occurred, the head and neck portion of the male continued its rhythmic bobbing over areas of empty substrate. The female's TFR was much reduced, and occurred during intervals sometimes spaced as far apart as 2-5 minutes. By 0005 h, the female lay partially coiled, though still linked to die male. Her TFR had ceased almost entirely, as had all movements of her body. The male's further efforts to entwine her were largely ignored. About 0020 h the female tried to crawl away to her burrow, diough still connected to the male. When he flowed along with her, still linked, continuing his chinning motions against her, she responded with vigorous tail vibration and a sudden lunge of her anterior body that resembled a partial strike. By 0040 h the pair had disconnected themselves. They retreated into separate hiding tunnels. Total copulation time: about 5 h. Male Combat Male combat was not very explicit, and much less violent or spectacular than breeding. It seemed to consist of little more than males coiling on top of each other at nightly intervals for several days prior to breeding. It was not always observed and seemed not to be a necessary requirement to trigger sexual activity of males (as witness the L. m. stenophrys breeding described below, which had only one 175

14 Figure 5. During courtship, the male Lachesis muta melanocephala flips over upside down on top of the tightly coiled female and commences a rapid back-and-forth sawing of his dorsum against hers. A "spare" female was coiled nearby during each L. m. melanocephala mating observed by the author.

15 male). Breeding never occurred on die days when male combatting was observed; rather it occurred on the days following. Presence of Spare Females In each instance of breeding of L. m. melanocephala a "spare," or additional, female was observed to be lying coiled beside the mating pairs (Figures 5 and 6). These females were superfluous to the actual copulation, but did not seem to be disturbed by it. The keeper inferred them to be prospective sexual partners to the male. Tight coiling of females prior to courtship seems part of the ritual activities of these snakes. The upside-down sawing activity of the male on the dorsum of the coiled female has been described. First Breeding of L. m. stenophrys, and How it Was Triggered The breeding of the single pair of L. m. stenophrys occurred in a substantiially smaller enclosure than the former breedings, this being 3.6 X 1 X 1 m high. Temperature gradients were maintained identically to the enclosure containing the L. m. melanocephala pairs. Humidity increase (to 95%) was accomplished through repeated wetting down of the mulch substrate with a pressurized sprayer. As no activity in these L. m. stenophrys had occurred despite temperature/humidity changes, the author hoped to trigger the breeding of L. m. stenophrys by using the chemical secretions left by the L. m. melanocephala pairs in the breeding enclosure. Removing the areas of wood mulch where residual secretions of the mating pairs were visible, die keeper transferred this matter into the smaller enclosure. The male L. m. stenophrys, upon locating this material, showed rapid TFR and bobbing head movements. He made frequent circuits of the cage area, returning to these areas and often coiling beside them, his head overhanging his coils. Any movement or vibration within the enclosure incurred a rapid TFR response in the male. The male repeated these activities for two nights. On 23 January, at approximately 1730 h, the female L. m. stenophrys emerged from her hiding tunnel out into the open gallery of the enclosure. She crawled over the male, chinning him at the nape and various parts of his dorsal body. As in the first breeding, however, the female gave some resistance once the male tried to intertwine her. This involved forcible tactics of the male to position her (upside-down "sawing," battering of his coils against her). Unlike the previously described breeding of L. m. melanocephala, the active motions of die male subsided sooner and the two snakes were observed to entwine to die full lengdi of their bodies, tails tightly wrapped. Expansion of the female's cloaca was achieved by the male physically bending die female's tail parallel to her body with his tail, which parted the female's vent area against his. This was attended by a vigorous grasping/twisting of the entwined tails. At length the female's cloacal region became engorged, the vent dilated to approximately 25 mm. About one hour passed before intromission was achieved. The lobes of the hemipenis were exposed and partially visible. Intromission continued until approximately 0010 h, when temperature was allowed to fall in the room. A milky residue was left on the substrate where penetration occurred. Duration of copulation: roughly 5 h. Both L. m. stenophrys returned to the same hide mnnel and remained coiled together for seven days. These specimens did not mate again. Conclusions Sexual activity in bushmasters is affected by temperature and humidity. It occurs widi a sudden increase of humidity after a period of increased heat and dry. Chemical placement is of importance in enabling males to find females. From the activities seen in this synthesized "rainforest" environment, the author conjectures that in the wild, the female bushmaster crawls a pattern area leaving chemical clues to her vicinity (the size of this area in the wild remains to be determined). The male locates the female through chemosensing this pattern. He may trail it to her whereabouts, or wait coiled nearby for her later return to die immediate area. Bodi male and female bushmasters use diese clues to rendezvous accordingly at a specific site described by placement. Female scent patterns may trigger sexual activity of odier females, as well as males. The bushmaster's middorsal ridge is employed in "sawing" movements against the female's dorsum in order to sexually stimulate her, and the male bushmaster inverts his whole body to do this, coiling upside down on the coiled female (Figure 6). As the male's dorsal ridge slides down into the space between the female's coils, this may have the effect of prizing her coils apart, which would enable him to sexually position her. "Defense" responses of the female, such as tail vibration, gular inflation, hissing and striking (without biting) may occur, during both courtship and copulation. Copulation takes place above ground in an open area, not in underground burrows or in hiding. All captive bushmasters here were observed to copulate outside their hiding tunnels, although these tunnels seemingly would be spacious enough to permit at least some aspects of courtship. A spare or prospective female may normally be present during copulation, coiled beside the mating pairs (observed in four instances). Multiple copulations may occur in both males and females. Copulations were observed involving female L. m. melanocephala 4, even after she had developed proximal swelling. During this period she continued to "scent" the cage floor just as before copulation. In L. m. melanocephala, courtship may begin diurnally, two or three hours before dark. Courtship of L. m. stenophrys (in the single instance observed) began at nightfall. Reproductively active females may cease feeding; males may continue to feed, although more sporadically. Males become sexually active at intervals of 5 to 10 days during the breeding season (assessed from multiple copulations of L. m. melanocephala). Duration of copulation is usually about 5 h (common to five instances observed here). Its termination may coincide with reduced temperatures of early morn- 177

16 ing hours, as would normally occur in the wild. Male combat consists principally of males coiling on top of each other at nightly intervals for several days prior to breeding. Breeding seems not to occur on the same night consecutive with combat, but on the following days. OVIPOSITION AND MATERNAL CARE OF EGGS Placement of Uric Acid Pellets As the pregnancy of the three female Lachesis advanced, they each deposited what appeared to be minute uric acid pellets (about the size of a dime) in a viscous fluid directly at the mouth of their simulated burrows. Such deposits had never been noticed before among nongravid bushmasters here. As the size of each deposit, and its visual composition, was consistent to each female, and placement before the opening of the burrow was always identical, the author inferred this to be related to the pregnancy. Studies by Jaeger and his associates have shown that male Plethodon cinereus deposit fecal pellets as territorial markers. The true function of these deposits of gravid bushmasters, if any beyond the obvious one of elimination, remains to be determined. Oviposition and Maternal Orientation to the Eggs Proximal swelling in all subject females was observed within six weeks of copulation. During this period, the snakes remained largely hidden, emerging only to take in what seemed inordinate amounts of water (above 200 ml). Female L. m. melanocephala 4 became very responsive to dismrbances in her environment. Unusual aggression was sometimes displayed toward her keeper. At times she would emerge from the back of her burrow (diis entailed crossing a distance of 2 m), and, advancing an additional distance of 3 m of open floor area, pursue him at close range. During these encounters her head was elevated to as much as 1 m above the ground. There was no tail rattling, body stiffening, or gular inflation, as is often observed in threat displays of this species. Rather her whole attention seemed directed toward the physical pursuit of the intruder. Here, her TFR was markedly different than at other times. The tongue was extended once quickly (for about 1 second), and withdrawn more slowly (taking about 3 seconds), giving the impression of a continuous flowing movement rather than a flick. The author has observed diis in other threat displays of this species, albeit uncommonly. When, inevitably, the keeper had been driven out of the enclosure, she went immediately back to her prospective nest site and coiled. Merely to bump the hiding tunnel was to provoke her to emerge again and "chase" her keeper. About 12 such incidents occurred over a 60-day period. As pregnancy advanced into its second month, the gravid females went through a period of several weeks of frequent and unusual crawling. This included crawling on their sides, and forcing their flanks against and between objects. This behavior seemed an attempt to move the eggs rearward. In the L. m. stenophrys female, the eggs were clearly visible through the body wall; her body seemed incongruously thick, and crawling appeared difficult. In both L. m. melanocephala females, however, the eggs were scarcely apparent. The L. m. melanocephala females seemed to grow thinner rather dian larger, a state perhaps compounded by their not eating. During the week of 10 April, L. m. melanocephala female 2 exhibited a marked inclination to hide. She moved back to the fardiest end of her hiding hinnel, and diere remained. Four days later, on 14 April, this snake was observed to coil in a peculiarly symmetrical way; her tendency was to stack her coils one on top of the other in levels, "beehive" fashion, her head balanced at the top where she faced outward. On 17 April, 101 days after copulation, she began depositing her eggs within the enclosing coils. There was no attempt to bury diem radier they were laid directly on the surface of the peat moss substrate within the tunnel. Her head was situated near the cloaca as die eggs emerged; she flicked her tongue at intervals against the egg surfaces as they appeared, and against her cloaca. The tail portion was slightly elevated, and moved as the coils shifted around the edges of the rising clutch. Occasionally the tail twitched from side to side as die eggs emerged. The eggs accumulated in the uniform circle of her coils, cementing one by one onto the clutch. The whole process took about 24 h. Her coils gradually rotated as the eggs were placed, but all die time die coils maintained their "beehive" symmetry of placement, stacked in two and three levels around her eggs, so that the eggs remained completely enclosed by coils. Upon examination, the eggs proved to be 12 in number, and just smaller than the eggs of large pythons (see Table 2). The female remained coiled around her eggs for about two days, at which time the author decided to remove her in order to artificially incubate them. It proved very difficult removing Table 2. Reproducrive data from 1993 captive breedings of Lachesis muta melanocephala and L. m. stenophrys. Xegg Number Copulation Oviposition Clutch dimensions Hatch hatched Adult pairs date date size (mm) date M:F L. m. melanocephala male 1 x female 2 male 1 x female 4 5 January Multiple dates February-March 17 April 9 May x63 86x June July 5:6 2:3 X neonate weight (g) X neonate total length (mm) L. m. stenophrys male 1 x female 2 23 January 4 May 16 76x July 5:

17 this female from her eggs. She tended to cling to them, repeatedly groping out for them with her coils to draw her body back around die clutch mass. Consequent tightening, and stiffening of her coils around the egg clutch, met every endeavor to lift her away. There was no tail rattling or defense postare, but some hissing occurred. Compared to normal behavior for this subspecies, she behaved in an amazingly docile way to her keeper. Unable to prize her free, the keeper allowed her to wind herself back around the eggs for a time. As she coiled around them again, she went through a period of flicking her tongue against the egg surfaces, her rostral bumping them lighfly. She seemed reorienting herself to dieir presence, and position. TFR was characterized by short but rapid modulations. Tongue extension differed markedly from what is observed both in courtship and preparatory to striking prey. In predation and courtship, TFR is characterized by long extension. In diis instance (and diroughout all aspects of egg guarding diat have been observed here), die tongue was but moderately extended, often just showing the tips of the forks, and moving lightly, almost tacmally, over the egg surfaces. In character it more strongly resembled the TFR that occurs after prey has been struck and is being identified preparatory to swallowing. Chiszar and Radcliffe (1976) reported diat ratflesnakes "did not begin to flick dieir tongues at a high rate until after they had struck prey, suggesting that the act of striking 'switches on' a chemosensory searching process which facilitates die location of dead or dying prey." Does oviparity "switch on" a similar process that enables the mother bushmaster to orient herself to her oviposition site, and remain there throughout the incubation period? The actions displayed by this female (and die odiers that were observed here) seemed at once tactual and chemosensory. As the mother snake could not be motivated away from her eggs, the eggs had to be lifted out from under her body. This had the effect of displacing her position on the peat moss. Once the eggs were removed, the female exhibited more evidence of chemosensory searching behavior. She explored the "empty" peat moss substrate widi her snout lightly bumping the surface. Her tongue movements were rapid, but shortly extended, the forks just passing the margin of the rostral scale. This continued for about 20 minutes, till again her head arrived at the spot where the eggs had been deposited. There she recreated die arrangement of stacked coils that had been observed previously. And so she remained, as though still brooding her now absent eggs. Upon removal, the eggs demonstrated a solid if still very soft and wet clutch with only two eggs not cemented to the mass. A third egg came away from the clutch as they were lifted out (this egg would later rot during incubation). All 12 eggs appeared to be viable. They were creamy white in color becoming brilliant white over die course of incubation. The eggs were transferred to a sealed 38-liter aquarium which contained a medium of vermiculite 1:1 by weight distilled water. Incubation was begun at 24 C. On 21 April, four days after the initial laying, a 13di egg was deposited by this same L. m. melanocephala female. This egg appeared viable. With carefiil attention to its original topbottom position, it was transferred to the vermiculite medium. This egg would later hatch on schedule with the others. On 9 May, L. m. melanocephala 4 laid nine eggs in another ninnel-burrow with a peat moss substrate. The eggs of this female were nearly identical to those of female L. m. melanocephala 2 in size, elongation and color. (See Table 2). The keeper left her brooding her eggs for a 3- to 4-day period before removing diem for incubation (Figure 7). Vigorous resistance to the removal of the eggs, including tail vibration and hissing, were noted of this specimen. Once eggs were removed, she was released again some distance from her nesting site. Here the female exhibited evidence of chemosensory searching. She explored the immediate area of enclosure, her head tracing a circular pattern on the mulch substrate. Tongue was but slighdy extended, with a high TFR (above 40 per minute). Once her head crossed the margin of the opening to die nesting place, TFR became markedly faster (above 60 per minute). Entering the tutmel, she paused at specific areas of the oviposition site, her snout bumping the moss substrate repeatedly. Here she raised her head from the substrate and directed it upward. For a 30-second interval she showed numerous slow fluid motions of die tongue, with long Figure 7. A female black-headed bushmaster {Lachesis muta melanocephala) with nine eggs. Photograph by the audior. Figure 8. A female Central American bushmaster (Lachesis muta stenophrys) with 15 eggs. Photograph by the author.

18 extension. At last, withdrawing herself entirely, she took her place as before, in a "beehive" arrangement of her coils, over the former site of the eggs. On 4 May, the female L. m. stenophrys, after some weeks of more than normally active crawling, deposited 15 eggs in the same arrangement of stacked coils (Figure 8). Again, it was 101 days since copulation, showing no difference between these two subspecies. The keeper again encountered extreme difficulty in separating this female from her eggs. She assumed typical bushmaster defense responses, including tail rattling and striking very unusual behavior for this long term captive L. m. stenophrys. Once prized off the eggs, she was moved to a new enclosure. This new environment was set up widi a burrow mnnel identical to the former, widi the exception, however, that the substrate did not contain egg scent. In her new location, the snake showed immediate signs of chemosensory searching. Short, rapid TFR was resumed, with moderate tongue extension, snout repeatedly bumping the substrate. The female explored the entire cage premises for a number of hours, as though attempting to trail her eggs by chemosensory means. Eventaally she went mto the burrow but came back out immediately. Unable to locate the egg scent anywhere in die enclosure, she focused her attentions on her own tail, and spent over 30 muiutes tongue-flickmg her own posterior ventral body, especially near the vent. The L. m. stenophrys eggs appeared distincdy different from the L. m. melanocephala eggs (Figure 9). They were of grayish yellow-white color, much less elongate and uniformly oval. From appearance alone one would not have immediately concluded them to be from the same species. A 16th egg, nonviable, was deposited one month later in an open area of the enclosure. The snake made no attempt to guard it. Duration of Egg Guarding and Arrested Feeding by Gravid Females L. m. melanocephala females 2 and 4, which were allowed to remain in die burrows filled with egg-scented peat moss, refused food throughout the 77-day period after the eggs were laid. They continued to guard, and wind about the absent eggs Figure 9. Eggs of Lachesis muta melanocephala (end tanks) and L. m. stenophrys (center tank). Note difference in shape elongate for L. m. melanocephala and oval forl. m. stenophrys. Color was significandy different also. Photograph by die author. at the farthest end of the tunnels. This mock brooding continued up until the last two weeks of incubation. The rigid "beehive" style stacking of coils was maintained at intervals diroughout diis time. These snakes, then, normally regular feeders, had refused food for a total of almost seven mondis. But even after food was at last taken, they remrned to their habitual nesting place in the hide tunnels, coiling in beehive fashion as before over die absent eggs. The L. m. stenophrys female, however, which had been quickly relocated to another enclosure where egg odor was not present, soon went back to regular feeding patterns and this within 10 days of egg laying. By contrast, L. m. melanocephala 2, once returned to her former nest tunnel, deposited a 13th fertile egg four days later; she continued to guard and brood diat spot for the duration of die 77-day incubation period, refusing all food until the approximate hatching date of the eggs. Thus it appears that the female bushmaster seldom leaves her eggs during die incubation period, even to feed. On but two occasions were these snakes observed even to drink (from bowls placed near the moudi of die burrows). Most remarkable is die female's capacity to fast while guarding her eggs she has already gone without food for three to four months during her pregnancy (noted of two L. m. melanocephala), and will continue to do so for another two to diree months until hatching is complete. Gravid L. m. melanocephala may grow extremely thin during this interval. L. m. melanocephala 2 and 4 lost approximately 15% of their overall mass while carrying eggs, totalling 30% after oviposition. Weight loss was evident even in the head region, which began to look visibly smaller. The spinal ridge was thus thrown into greater prominence, causing them to look emaciated and sickly. While collecting bushmasters in western Surinam, the author had often noted how male specimens were nearly always larger ttian females, usually about cm longer, but always of greater mass even when of equal length. This is explicable when one takes into account how the female may fast for upwards of six months out of each breeding year. In the generally cool temperatures of rainforest conditions, her reduced metabolism may play a role in helping her through this long period of inactivity and fasting. Conclusions The bushmaster's maternal/protective response to her eggs is highly developed. Bushmasters are oriented to their eggs by chemical clues, and will guard and "brood" an area even where eggs are no longer present, providing egg scent is still available. Removing the female to a novel environment where egg scent is not present will trigger a chemosensory searching process involving exploratory trailing to relocate her eggs. In the absence of chemical cueing, the snake responds with appropriate exploratory activities to relocate it. Apparently, once the maternal egg guarding process is "switched on" it is not immediately "switched off by deprivation of chemical cues from the eggs. Female bushmasters, so long as egg scent is present, fast for the duration of the incubation period. The presence of the egg odors not only "switches on" the bushmaster's vomerona- 180

19 sal system, but triggers a response that inhibits almost all other functions (including feeding) but those related to egg guarding. When the relevant chemical cues on the substrate are removed, the snake returns to normal feeding patterns after a few days (in the case of the L. m. stenophrys female, about 10 days). Such occurrences suggest a response-sequence integration mechanism such as been recorded of rattlesnake feeding episodes (Chiszar et al., 1977). In this case, a response (oviposition) puts the animal into a position to detect stimuli (egg odors) which release the next response of the sequence (egg guarding and orientation to their location). How does die bushmaster know when to cease guarding her eggs and begin normal patterns of predation? Removal of significant chemical stimuli may cause these reptiles to revert to former responses, which may include detection of other stimuli such as those chemical ones emitted by prey. Not only do egg-guarding female bushmasters cease feeding during the period while they guard the eggs, they seldom leave the eggs even to drink water. The snake seems to enter a state of semitorpor during the period of egg guarding, her activity consisting of little more than the shifting of the placement of coils. Female bushmasters may defend their eggs vigorously, or they may not but they strongly resist any effort to be taken from them, and make efforts to cover diem (i.e., protect them) with their coils. They do not abandon their eggs at any time after oviposition, but remain oriented to them in a protective manner throughout the entire 60- to 77-day period of the incubation process until the eggs have hatched. This behavior does not differ in the wild. As a means of lending support to this last statement, the audior cites a field experience occurring while collecting Lachesis muta muta in Soudi America: In July 1984 die author was called by local hunters to excavate a paca (Agouti paca) burrow in West Surinam (Apoera, Dist. of Sipaliwini) where a bushmaster was diought to be present. The hunters, using dogs, had trailed a tabba (paca) to its suspected burrow in an area of primary forest, on a shallow hillside. Upon sounding the depth of the tunnel with a long stick, the hunters had encountered a buzzing noise which they suspected might come from a malcasneki (bushmaster) hidden in the burrow also. The author, carefiiuy digging out a portion of the hillside, found a 2.4 m female bushmaster coiled around her eggs in the paca burrow. These eggs, 19 in number, were removed, and incubated artificially. Eight of them hatched fifteen days later; demonstrating that, from die known incubation time of days, this female had already brooded her eggs for at least days prior to discovering her. So far as is known, no confirmed reports exist in literature of the finding of bushmaster eggs postparmm, but it is apparently not an uncommon occurrence for hunters in some parts of Central and South America. The author has extracted 11 bushmasters of three subspecies from paca burrows, and in a number of instances live pacas and in one case an armadillo (family Dasypodidae) were present in the burrow channel as well. It is worth noting that the L. m. melanocephala specimens of this study were, with one exception, all collected through excavating paca burrows. On one occasion one male and two female adult L. m. melanocephala were taken from the same burrow (Figure 10). From the above instances cited, it is possible to conclude that bushmasters do indeed show maternal care for their eggs in the wild. Once oviposition takes place, bushmasters do not abandon their eggs, but guard them for the full duration of the incubation period. The likely location for oviposition is in a secure, dark and humid burrow underground. Below ground areas like the paca burrow make for stable temperature and humidity. If the burrow selected is in a hillside (a typical site) both eggs and snake will escape flooding a constant danger for eggs during die rainy season. As die eggs are not buried in wood humus or debris (as was formerly believed), but lie exposed on the burrow floor, maternal guarding may be an important evolutionary response against predators. Additionally, the snake is able to ward off inadvertent destruction of her eggs by the active mammal burrowers that have not only constructed her lodging and nest, but threaten them with their persistent presence. Thus, egg guarding is of survival importance to diis species. Ontogenetic Color and Pattern Changes, and Relative Sizes of Neonates Bushmaster neonates emerge with very faint, indistinct Figure 10. A paca (Agoutipaca) burrow on Peninsula de Osa, Costa Rica, out of which one male and two female adult Lachesis muta melanocephala were extracted. The head of one snake can be seen protruding from the opening. The two men are local paca hunters who informed the author after discovering the snakes. Photograph by the audior. 181

20 markings. They do not begin to demonstrate adult coloration until after a year or more of age. Juvenile specimens of L. m. stenophrys (Figure 11), even up to flieir second year, may lack black pigmentation in the dorsal markings. In hatchlings the pattern is only marginally darker dian die ground color, and only the postocular stripe exhibits die overall melanism later seen in the rhomboidal markings of adults. The general ground color of L. m. stenophrys neonates is orange. The iris of young and neonate L. m. stenophrys and L. m. melanocephala is red-brown, becoming obscure and heavily pigmented as the animal grows, by the first year black within the postocular stripe. The ground color of L. m. melanocephala neonates (Figure 12) is yellowish tan, becoming more strongly yellow, and lighter, as they mature. The heads of L. m. melanocephala are uniformly dark dorsally like the parents, but with the addition of a faint line of color separating the postocular stripe from die melanistic dorsum of the head. This ontogenetic trait of neonates is recorded in the original description of L. m. melanocephala (Solorzano and Cerdas, 1986). It fades gradually over the course of development (by age three months the line has begun to fade perceptibly, by six mondis it is no longer visible in the increasingly pigmented dorsum of die head). Adult specimens retain vestiges of this juvenile trait in die form of a divisionary color line below the angle of the jaw. In some adult specimens diis color line is still visible in the black head cap. The melanistic head dorsum of L. m. melanocephala is therefore not die result of a union of the postocular stripes across the dorsum of the head, but rather an independent and unconnected blotch simated there, diat gradually blackens over the first months of life. The "braided" lepidosis (scales outlined with darkly pigmented underskin) typical of adult L. m. melanocephala is also present in neonates, aldiough somewhat less defined. It is more evident posteriorly. Like die black head dorsum, diis trait seems distinct from other Lachesis. Like many neonate pit vipers, bushmaster newborns possess a bright orange or yellowish tail tip, presumably useful in attracting mammalian insectivores as prey items. Young bushmasters have proportionately much larger heads than adults, visibly distinct on their relatively slender bodies. Neonate L. m. melanocephala, with a total length (TL) of 533 mm at hatching, have heads fully 31 mm long by 19 mm across. This large head size enables them to accept a wide range of prey items. Neonate and juvenile bushmasters swallow proportionately much larger prey than adults. At hatching neonates are already capable of feeding on adult mice, and even small rats. The hatchlings displayed marked defense reactions upon encountering their keeper for the first time. This included audible tail vibration, and striking from an elevated head stance, even as they emerged from the egg. Specimens of L. m. melanocephala struck repeatedly, and advanced bodily to do so. A total lengdi (TL) of 1180 mm, and a weight of 675 grams, was attained by neonates of the first group of L. m. melanocephala by January 1994 a growth rate of over 100% in six months. L. m. stenophrys achieved a TL of 1070 mm, and a weight of 450 grams after the same period of time. Out of 28 hatchlings there have been no mortalities thus far. This is the largest captive reproduction of Lachesis to date, exceeding in one year die Dallas Zoo reproductions of L. m. muta irom 1987 to Aclinowledgments The audior wishes to diank Theo Henzen of Paramaribo, Surinam, for invaluable experiences in South America, and the hundreds of local hunters and Indians through the years who made finding bushmasters possible, and the jungle livable, in Brazil, Surinam, Guyana, Ecuador, Panama and Costa Rica. Figure 11. Neonate Lachesis muta stenophrys. The only black on diis animal is in die postocular stripe, a condition that persists until the snake's second year, when the dorsal markings become progressively darker and more contrasting against the ground color. By six months of age the iris will have turned uniformly black. The tail tip is bright orange and may act as a lure to insectivorous mammals. Photograph by the auuior. Figure 12. Neonate Lachesis muta melanocephala. Note the presence of a light color line delineating the postocular stripe and separating it from the dark head cap. This is an ontogenetic trait dial gradually fades away as the snake reaches its first year, and the dorsocephalic region becomes uniformly black. Apart from the postocular stripe, there is no true black pigmentation in neonate L. m. melanocephala, or in neonates of any bushmaster subspecies. In L. m. melanocephala, the head cap turns gradually black over the first year of the snake's life, and by adulthood will be blue-black. The ground color will become a very light straw-yellow ranging to cream white, which contrasts sharply to the black dorsal markings. Photograph by die audior. 182

21 Thanks to Maria de Fatima, Wilson Fernandes and Guisseppe Puerto for expert help at Instituto Butantan, Sao Paulo, Brazil; Alma Hoge for important assistance and gracious company; Don Boyer of Dallas Zoo; Don Wheeler and Mike Parker for illustrations; William S. Burroughs for much-needed encouragement through the years. This project was funded by the free enterprise system, paid for by the sale of reptiles both venomous and not venomous to zoos and private individuals. The intent of this study was to make profit through the reproduction of a rare and increasingly endangered species; the information generated here is a byproduct of that process. In light of his success, die author wishes to acknowledge a capitalistic society where it is still marginally possible to pursue life, liberty and happiness without die approval of self-styled authorities in government offices. Lastly, it should be mentioned that studies such as this, by a private individual, would not be possible in states where the keeping of venomous snakes is prohibited, where mob opinion is allowed to prevail over individual judgement, where equal rights between individuals and special groups do not exist, to the extent that only "qualifying" institutions (the institutions write the rules on who qualifies) are allowed to keep many forms of venomous reptiles. The author wishes to acknowledge the state of North Carolina for respecting man's animal right to coexist widi other animals of his own choice, and for not seeking to limit and deform the human embracing of nature. References and Literature Cited Amaral, A. do On die oviparity of Lachesis muta Daudin Copeia 1925: Bolaiios, R Serpientes venenos y ofidismo en Centroamerica. San Jose: Editorial Universidad de Costa Rica. Boyer, D. M., L. A. Mitchell and J. B. Murphy Reproduction and husbandry of the bushmaster (Lachesis m. muta) at the Dallas Zoo. International Zoo Yearbook 28: Boyer, D. M The bushmaster program at the Dallas Zoo: husbandry, reproduction and research (flyer). Department of Herpetology, Dallas Zoo. Burchfield, P The bushmaster Lactow WKto in captivity. Internafional Zoo Yearbook 15: Campbell, J. A., and W. W. Lamar The venomous repfiles of Latin America. Ithaca, NY: Cornell University Press. Chiszar, D., and C. W. Radcliffe Rate of tongue flicking by rattlesnakes during successive stages of feeding on rodent prey. Bull. Psychon. Soc. 7: Chiszar, D., C. W. Radcliffe and K. M. Scudder Analysis of the behavioral sequence emitted by ratflesnakes during feeding episodes. I. Striking and chemosensory searching. Behav. Biol. 21: Chiszar, D., J. B. Murphy, C. W. Radcliffe and H. M. Smidi Bushmaster (Lachesis tttuta) predatory behavior at Dallas Zoo and San Diego Zoo. Bull. Psychon. Soc. 27: Ditmars, R. L Snakes of die world. New York: Macmillan The making of a scientist. New York: Macmillan. Greene, H. W., and M. A. Santana Field studies of hunting behavior of bushmasters. American Zoologist 23:897. Hoge, A. R., and A. R. Lancini Sinopsis de las serpientes venenosas de Venezuela. Publ. Ocas. Mus. Cienc. Nat. Caracas, Zoologia 1:1-24. Hoge, A. R., and S. A. R. W. D. L. Romano. 1976/77. Lachesis muta rhombeata [Serpentes: Viperidae, Crotalinae]. Mem. Inst. Butantan 40/43: Jaeger, R. G., J. M. Goy, M. Tarver and C. E. Marquez Salamander territoriality: pheromonal markers as advertisement by males. Animal Behavior 34: Jaeger, R. G., and E. A. Home Territorial pheromones of female red-backed salamanders. Ethology 78: Martinez, V., and R. Bolaiios The bushmaster, Lachesis muta muta (Linnaeus) [Ophidia: Viperidae] in Panama. Rev. Biol. Tropical 30: Ripa, D The reproduction of the Central American bushmasters (Lachesis muta melanocephala and Lachesis muta stenophrys) for the first time in captivity. Vivarium 5(5): Solorzano, A., and L. Cerdas A new subspecies of the bushmaster, Lachesis muta, from southeastern Costa Rica. J. Herpetology 20(3): Vial, J. L., and J. M. Jimenez-Porras The ecogeography of die bushmaster, Lachesis muta, in Central America. American Midland Namralist 78:

22 184 Bull. Chicago Herp. Soc. 29(8): , 1994 Absolutely prohibited After years of exploitation, including the taking of newly hatched babies for varnished paperweights and the adults for turtle soup, alligator snappers are protected throughout their historic range. The species is listed as Category 2 under die U.S. Endangered Species Act. State protection will have to suffice until die species' status is reviewed by the U.S. Fish and Wildlife Service in [Wildlife Consen/ation May/ June 1994, from Karen Purnweger] New rules published Jim Harding sent along a copy of Order #DFI from the Director of the Michigan Department of Namral Resources, which states: "I hereby order: That it shall be unlawfiil to kill, take, trap, possess, buy, sell, offer to buy or sell, barter, or attempt to take, trap, possess or barter any reptile or amphibian from the wild, or the eggs of any reptile or amphibian from the wild, except as provided within this order." Certain species are prohibited utterly without special permit from that office, including: eastern massasaugas, Blanding's turtles, wood mrtles, spotted turtles, eastern box mrtles, black rat snakes, cricket frogs and any reptiles and amphibians protected under the Endangered Species Act. Commercial take is limited to snapping hirtles with a carapace length of 12 inches or more and green frogs subject to legal limitations. "All reptiles and amphibians taken for personal take shall not be bought, sold or offered for sale." There are possession limits for all legally collected animals. For a full copy of the act, write: DNR, Box 30028, Lansing, Ml Don't say you didn't know it looks as if the state means business on diis one. Box turtle update Those who attended a recent CHS meeting were made aware of a grass roots movement from the New York Turtle and Tortoise Society (including CHS member and regular contributor to this column, Alan Salzberg) to have the eastern box turtle immediately listed on Appendix III of the CITES treaty. In addition, they hope to have its statos changed to Appendix II at the next CITES meeting in November. Why? Well, as reported in this column previously, more than 30,000 box mrtles were legally shipped overseas last year, and the species is becoming uncommon in many parts of its range. Pressure on the species began to build once Mediterranean tortoises were no longer legal for sale in nordiern Europe. Animal dealers switched to box turtles instead. Collectors here are paid about a dollar, and the animals are sold in Europe for up to $200 (according to a recent houseguest). However, die mortality of die transshipped reptiles is very high. CHS member Jim Harding suggested that it may be 90 percent within diree years. He added, "Recent research has clearly shown that such species have virmally no harvestable population surpluses, and cannot be harvested on a sustained basis." [Lansing, MI State Journal May 25, 1994, from Jim Harding] The old shell game A town near London, Ontario, Canada, has sponsored turtle HerPET-POURRI by Ellin Beltz races for 20 years, but diis year the Ministry of Natoral Resources requested that die organizers offer instructional seminars along with the races. It seems that permits are required to catch midland painted mrtles in the province, and that special dispensation was given diose collecting for the races. All the turtles will be released after die event. [London, Ontario, Free Press July 15, 1994, from Ted Teachout] Shell shocked, but healing A yellow-bellied slider was found in Apex, Nordi Carolina, with a 3 '/z-inch gash in her 10-inch-wide shell, which exposed her spinal cord. Vets at the Nordi Carolina State University's College of Veterinary Medicine said that the cut looked as diough it had been caused by a shovel or a golf club. A team of doctors washed die wound, and injected die turtle with antibiotics, then built a temporary shell to cover the gash. The mrtle was released into a pond outside Raleigh. [Baton Rouge, LA Advocate June 30, 1994, from Ernie Liner] Mosqulto/toad compromise reached Efforts to control mosquitos by pesticide spraying can continue in areas of Albany County, Wyoming, where Wyoming toads are not found, under a compromise accepted by the U.S. Environmental Protection Agency (EPA). In 1992, the EPA angered local residents and ranchers by banning the use of 43 pesticides in a 970-square-mile area. A toad search will be conducted and areas which show as wetlands on aerial photographs will be examined for the vanishing amphibian. After an area has been certified "toad-free," pesticides can be used for mosquito abatement. [Casper, WY Star-Tribune April 16, 1994, from J.N. Stuart] Fangs for the mammaries? The Arizona Republic (June 14, 1994, from Tom Taylor) reports "At a west-side topless bar, lounge lizards aren't the only reptiles lurking in the shadows. Six rattlesnakes have joined the cast... a guaranteed conversation starter. The fanged serpents, kept in a special cage, are popular with the patrons, but not widi state Game and Fish Department officials who say ratdesnakes can be displayed only for educational purposes.... The snakes are kept in a specially designed terrarium, which consists of a 750-gallon tank inside a 1,000-gallon tank, with sophisticated environmental controls, a security system and authentic desert flora.... The cage has soundproofing... the snakes seem to ignore die topless dancing altogether." Newts in space Four Japanese red-bellied newts, quickly dubbed "astronewts" by die bright boys at NASA, were loaded onto the space shuttle Columbia after beating all comers in an egglaying competition. Besides the newts and a group of Japanese fish called medaka, die nonhuman passenger list includes six goldfish, 126 jellyfish, 144 newt eggs, 340 medaka eggs, 180 toad eggs and six toad testes for the purposes of fertilization, 11,200 baby sea urchins and 500 flies. A control group of die

23 same kinds and number of animals will remain on eardi and undergo die same experiments as the space critters. Scientists are interested in examining how eggs and animals develop and behave in weightlessness. [Chicago Tribune July 8, 1994] Recent television reports suggest that one or more of the newts died in orbit. If anyone fmds the rest of this story, I'd really like to see it. Headstarting controversy explained Jean Karnes sent a copy of the Summer 1994 Marine Conservation News from the Center for Marine Conservation, written by Deborah Crouse, which expresses ttieir point of view on the closing of the Galveston Head Start Center for Kemp's Ridley Turtles. [See also Dr. Grouse's letter in die March 1994 "Tympanum," Bull. Chicago Herp. Soc. 29(3) ] The Galveston center was supported for many years by die efforts of H.E.A.R.T., whose founder and tireless leader, Carole Allen once spoke to the CHS. Apparently hearings were held in March and "five witnesses specifically noted that 'headstarting' Kemp's Ridley sea turtles could not be substituted for Turde Excluder Device (TED) requirements in the shrimp fishery." Louisiana Representative Billy Tauzin chaired the hearing. Rep. Tauzin has long represented the interests of shrimpers and opposed TEDs. Now he is championing the raising and releasing of baby sea turtles "so diat shrimpers won't have to use TEDs." The National Marine Fisheries Service (NMFS) stopped headstarting after 15 years after a review of the project by outside scientists indicated diat they did not feel that the headstarted turtles survived to reproduce. Until data indicating the 16,500 hatchlings already released are surviving/reproducing, it is unlikely that die project will be refunded. In all fairness, it should be pointed out that the project has not been going on long enough for any of its hatchlings to have become reproductively active. In addition, protection of adults and nestlings on die beaches at Rancho Nuevo, Mexico, may in the long run have more to do with the potential survival of the species in the wild. Crouse concludes: "To rely on headstarting as an alternative to TEDs would waste hundreds of thousands of taxpayers' dollars, raising turtles simply to be caught and drowned in shrimp trawls soon after release. As every single witness, even those in favor of restarting headstarting, recognized at die hearing, captive propagation or rearing stands no chance of helping an endangered species if the original threats are not addressed as well." Snake tales National Wildlife (April/May, 1994, from Mark Witwer) has a great article about Puerto Rican boas diat eat fruit bats. I suggest that you check it out in your local library because there is no way I could do justice to the photos, and the article is well written, too. Snakes "set up house" in an attic in a community near New Orleans according to a Times-Picayune report (May 21, 1994, from Ernie Liner). The family had diought diey had rats. It turned out to be rat snakes. One said, "I haven't slept in three days," Two snakes were captured, up to four more have been spotted. She added "They make a nasty sound, sickening, I turn the TV and radio up, but I still could hear it." A "Rattlesnake Show" was held at die Stockton, Illinois, Business Expo. "Snake experts Dan, John and Joe Roth... took the stage with a variety of snakes, including Western and Eastern Diamondback and Timber rattlesnakes, a copperhead snake, albino and regular reticulated Burmese pythons [sic], a corn snake and a bull snake." The show was reportedly educational and informative. The article continued, "The Rodis participate in die yearly World's Largest Rattlesnake Hunt in Sweetwater, Texas, where they both collect and butcher rattlesnakes.... Roth keeps his 30 Western Diamondbacks, two Eastern Diamondbacks, one Timber and one Rock rattler and his copperhead in his home." [Freeport, IL Journal-Standard April 2, 1994, from Lester Telkamp] CHS member Rick Dowling was recently feanired in his local Prattville, Alabama, paper, Prattville Progress, which he kindly sent along. Seems that since 1988, he has advertised in that journal offering to come and get snakes which are bothering people on their property. He said, "The first year or so, the only calls that I got were after people had killed die snake and wanted me to come out and identify it. But now I get about two calls a week from people who want me to relocate the snakes.... I came across a man who was shooting at several snakes in a small creek. He was convinced the snakes were water moccasins. I told him that they were simple water snakes, but apparently his father and his grandfather before him had decided that all water snakes were water moccasins. 1 reached down and picked up one of the snakes, just to show him that it was harmless and it bh me. Well, die man just freaked out and told me that I was going to die. As you can see, I'm still here." Let's see if I can sort this one out... A man who owned a "surly" 10-foot python asked a friend to feed it for him. The feeder bought a small brown and white rabbit for $21.49 and took it to work widi him. A co-worker flipped out that die bunny was snake food and (as the police report reads) "took the rabbit and brought it to her vehicle. The complamant [who had bought die rabbit] went out to [the co-worker's] vehicle, but she refiised to open the door to give him back the rabbit... [she] then drove off and took his rabbit. The complainant stated he contacted [the co-worker] by telephone because he thought it was a joke... [she] refused to remrn the rabbit because she did not approve of the use for the rabbit." Since then, the feeder bought another rabbit, and was fired from his job. He believes it was due in part to negative publicity surrounding diis incident, aldiough an official at die company told die reporter die bunny heist had no effect on die feeder's employability. [Daily Herald (north suburban Chicago), April 25, 1994, from Clover Krajicek] "Psycho on the loose in Mokena!" screamed a recent headline on Herald-News (Joliet, IL, from K. S. Mierzwa). The "Psycho" of die headline is a snake which escaped while visiting the shop in which his human worked. Seemed the human (stop me if this gets too familiar) left die snake in his truck with the window cracked one inch. Somehow, an 8'/4-foot Burmese python that is reportedly six inches around squeezed out of the truck and left. The owner said, "I got it for an 185

24 investment. I was going to breed them.... He's worth some big money." [At the risk of having more people cancel their memberships, I'd like to propose that the snake isn't the only psycho in Joliet. EB] Where does a 1000-lb crocodilian sleep? In this case, in traffic. A 12-foot alligator apparently passed out while crossing a highway, causing a 1'A-hour traffic jam while officers tried to wake it up. Some artist ought to do a poster on how to resuscitate an alligator. The police tried everything. One put his squad car right in front of the critter and set off the siren and blue lights. Anodier poked at it with a stick. The fire department showed up. When it finally did wake up, one police officer said, "He had an attitude problem. A gator that size does what he wants to and goes where he wants to go. I was open for suggestions." [Houma, LA Courier June 16, 1994, from Ernie Liner] Thanlts to everyone who contributed this month and to J. H. Schoenfelder, Kathy Bricker, Clover Krajicek, Ray Boldt, J. N. Smart, and Marty Wnek who sent smff I enjoyed reading but didn't use. You can become a contributor, too. Send clippings with publication/date slug and your name firmly attached with tape (or die whole shebang photocopied, please, please) to: Ellin Beltz, 1647 N. Clybourn Avenue, Chicago, IL Bull. Chicago Herp. Soc. 29(8): , 1994 In ray January and March columns I wrote about the proliferation of new sources of herp-related information and gave a few examples. Since then I have found more such interesting and potentially useful information. For those of you who might not have seen these booklets or magazines, the following is a short summary. The Herpetocultoral Library series contains a booklet entided The General Care and Maintenance of Savannah Monitors and Other Popular Monitor Species by Michael Balsai. It offers several suggestions that I particularly wanted to highlight (I found die booklet very good). Under the heading "Acclimation" it mentions that some herpetoculturists advocate giving a newly acquired animal an electrolyte solution like Gatorade instead of water for the first couple of weeks. I have seen diis advice elsewhere, and to me it makes good sense. An animal that is possibly (probably) very stressed and dehydrated might have its electrolytes diminished or out of balance. Under "Maintenance" Balsai writes that a cage should be cleaned with some sort of mild disinfectant at least once a week. He stresses not to use phenolic compounds such as Lysol*, as the residues could be toxic. He recommends either a dilute solution of Nolvolsan-S (chlorihexidine) or dilute sodium hypochlorite bleach (e.g., a 5% solution of Clorox*). Under "Feeding" he warns of the risk of Salmonella infection if you feed raw eggs or raw chicken to your animal. Both should be cooked first. He also points out that feeding nonembryonated (infertile) eggs can cause a biotin (vitamin B4) deficiency. This is because the whites of nonembryonated eggs contain avidin, which in excess can cause this deficiency. Embryonated (fertile) eggs can be obtained in many healdi food stores. Another paragraph in this same section calls attention to a recommendation of Frye (1981, 1991) that prey items should be offered as food soon after removal from their rearing cages. Helpful Herp Hints by Dermis Engler This is because the residual food in the prey's digestive tract can help supply the animal being fed with vitamins, especially vitamin C. Vitamin C from this source may aid in preventing stomatitis (mouth rot) and maintaining skin integrity. The section on "Grooming" discusses claw trimming. A brief oudine of the procedure: 1.) Use a cotton swab or gauze soaked in alcohol or Betadine* to disinfect the claws and surrounding areas. 2.) Use a nail clipper or cutter, then smooth away any sharp edges with a nail file or emery board. 3.) If you accidentally nick the blood vessel, wipe die claw with some disinfectant and apply cornstarch or KwikStop. An alternative is to let a veterinarian do it for you. Also, some pet stores that deal heavily in birds have personnel who can do this for you; they have to trim the claws of the large parrots regularly. One last item from this booklet: under "Nutritional Disorders" is a paragraph on metabolic bone disease / calcium deficiency. This can be a problem in young monitors fed exclusively on crickets or pinky mice not supplemented with vitamins and minerals. Symptoms could include "soft" jaws and/ or back and limb deformities in juveniles and swollen "smooth" hind limbs and/or swollen or deformed lower jaws in adults. One way to avoid this disorder in baby monitors is to get them to eat canned cat or dog food once or twice per month. This should supply the vitamins and minerals your monitor requires. The author recommends the poultry flavors, which are somewhat more namral and less fatty. The following hints are excerpted from three separate issues of Reptiles magazine. An article on amphibians in the October 1993 issue mentions that they do not take well to handling. The "slimy" protective coating is damaged by human touch. This can lead to the animal drying out or being poisoned by the salts or oils from human hands. Also, some species have toxins that may cause some discomfort to die handler. If you must handle them, wash your hands before and after. Washing first and leaving your hands wet makes for 186

25 less chance of damaging die animal. Washmg after handling minimizes your risk of transferring toxins to your skin, mouth or eyes! Frogs and toads have a tendency to empty their bladders to discourage handling, so always hold the back end away from your eyes. Watch your cats and dogs: diey have been known to die from just one bite on certain amphibians. In the same issue Bill Love gives tips to accelerate the growth of young animals: feed often, feed smaller food items, provide a heat gradient for your animal, handle the animal as litde as possible, and feed thawed, previously frozen food. The freezing ruptures (he cell walls of the food animal making it easier to digest. This is also a good idea when feeding weak or sick animals. Bill does caution that accelerated growth might shorten an animal's life. In the December 1993 issue, Roy Pinney mentions using a "flexible claw pick-up," also known as a "nut retriever" to retrieve uneaten food or whatever from a cage. This will cause less disturbance to the animal and might also benefit you if you have a particularly aggressive animal. The device is available in 15- and 24-inch sizes at hardware and auto accessory stores. In die same issue a formula is given on how to datermine how much to feed an animal. The formula for a reptile is Kcals = 10 X (body weight in kilograms)"''. Without boring most of you with the details (those who want them can read the article), it's figured out from this that a one-pound snake can get enough calories to meet its metabolic requirements by eating a 50-gram mouse every 15 days. This is greatly oversimplified because odier factors such as age, healfli, pregnancy, environment, stress and species must also be taken into account. Furthermore, life in captivity differs from life in the wild, where a lot more energy is expended m fmding and subduuig prey. Also, prey items in the wild are probably not so well fed as prey items in captivity. The thrust of diis article is not how little you can feed an animal but that you should not let your animal get obese. This can shorten the life span of your animal. In the August 1994 Reptiles, I read the following very disturbing piece by Roy Pmney (at least to me it was): Action Alert Seventy-one thousand box turdes were exported from die United States during die past three years for the overseas pet trade! North American box turdes (Terrapene spp.) are already under severe threat due to habitat loss and degradation, and wild populations are decreasing dramatically. The New York Turtie and Tortoise Society, die Turde Recovery Program, die New York Zoological Society and odier organizations across die country are seeking to control this continuing harvest through CITES (Convention on International Trade in Endangered Species) regulation. Writing to Dr. Charles W. Dane, U.S. Fish and Wildlife Service, Washington DC 20240, to support such legislation may make a big difference. I personally urge you to write to Dr. Charles Dane every letter helps. Also, I ask you to keep those hints coming in. If you would like to share a bit of knowledge that has helped you in taking care of herps, send it to: Chicago Herpetological Society, Helpfiil Herp Hints, 2001 N. Clark Street, Chicago IL Happy Herping! Literature Cited Frye, F. L Biomedical and surgical aspects of captive reptile husbandry. Edwardsville, KS: Veterinary Medicine Publishmg Company Biomedical and surgical aspects of captive reptile husbandry, 2nd edition, 2 vols. Malabar, FL: Krieger Publishmg Company. [Also published by T.F.H., Nepmne City, NJ, under the tide Reptile Care: An Atlas of Diseases and Treatments.]

26 Bull. Chicago Herp. Soc. 29(8):188, 1994 A LETTER FROM THE CHS PRESIDENT If you read the upcoming events section of die herp magazines and newsletters, you may have noticed the announcement for the Second World Congress of Herpetology last year. Many of you may have wondered about this organization and its purposes information that may not be readily available to the hobbyist because die congress was attended mainly by institutional herpetologists. I will give you some background, as some issues that affect all of us were addressed at the Second World Congress. The World Congress of Herpetology was formed in 1982 at a joint meeting of die Society for the Smdy of Amphibians and Reptiles and the Herpetologists' League, a meeting that also included representatives of the world's odier principal herpetological societies. The purpose of the World Congress was to establish a global forum where herpetologists from around the world could meet regularly, discuss their ideas and present their findings. To this end, the First World Congress of Herpetology was held in 1990 at the University of Kent in Canterbury, England. Lecmres were given, papers were presented, herpetologists and smdents got together in the pubs and exchanged ideas. The Second World Congress of Herpetology was held last December in Adelaide, Australia. I was formnate enough to attend this event. While there, I was asked to participate in drafting a resolution which would recognize die importance of amateurs in the field of herpetology. Now I realize that many of us do not consider ourselves amateurs and may even be offended by the use of the term amateur to describe people who, although lacking a degree in biology or herpetology, are quite expert in die husbandry of reptiles and amphibians. Many "amateurs" are so successful at keeping and breeding herps that they are able to support themselves and dieir families by such "amateurish" activities. The field of herpetology has broadened from scientific smdy to include husbandry, conservation and commercial management of reptiles and amphibians. This new group of people and die value of their contributions to the field of herpetology must be acknowledged regardless of their academic standing. The amateur support resolution adopted by die Second World Congress is the official recognition by the worldwide community of herpetologists of the importance of the contributions made by amateurs to the field of herpetology. Many herpetologists became interested in their science by collecting and keeping reptiles and amphibians as children. There are those who never received formal training and maintained a lifelong dedication to swdying these animals. While we can arbitrarily define what is a professional herpetologist and what is an amateur herpetologist, die line between die two cannot be drawn when we compare knowledge, commitment and contributions. The amateur support resolution levels the The Tympanum playing field. Currentiy, laws exist in the U.S. and many other countries discriminating among amateur and institutional herpetologists and zookeepers. These laws govern who may obtain collecting permits, participate in field sttidies or keep an animal in captivity. It is to be hoped that this resolution passed by the Second World Congress will serve to end die discrimination and allow all knowledge and contributions to be viewed equally and objectively, regardless of the source. Resolution by the Second World Congress of Herpetology 1. RECOGNIZING thatamateurherpetologistshavecontributed substantially to the body of fundamental knowledge which underlies herpetological research and further diat diey contribute to die knowledge needed to effectively manage and conserve wild populations of reptiles and amphibians, and 2. ACKNOWLEDGING tiiat many herpetologists develop tiieir interest and expertise with captive amphibians and reptiles, 3. AND further acknowledging that the collection of many species of reptiles and amphibians from the wild by amateur herpetologists, zookeepers and instimtional herpetologists is a negligible factor in the decline of their populations, and 4. CONCERNED that many current conservation laws are ineffective for the conservation of biodiversity in that: a) they are written for ease of enforcement b) they seriously inhibit and restrict herpetologists from pursuing their responsible interests and from contributing to the body of information tiiat is essential for die effective conservation of reptiles and amphibians THE SECOND WORLD CONGRESS OF HERPETOLOGY at its meeting in Adelaide, Australia, from 26 December 1993 to 6 January 1994: 1. Encourages continuing and wider amateur involvement in developing herpetological knowledge, and 2. Urges review of current legislation and its administration a) to ensure diis effective conservation of the biota, and, b) to support contributions by amateurs to herpetology. Moved: Harald Ehmann, Australia Klaus Henle, Germany Marcia Rybak, USA Seconded: Chris Banks, Australia Glenn Shea, Australia Gerry Swan, Australia Adopted: In favor: Against: 1 Abstentions:

27 Bull. Chicago Herp. Soc. 29(8): Photo Essay: 1994 CHS Expo Photography by Tony Rattin The scenes below are from the first CHS Live Amphibian and Reptile Expo, held June 4-5 at Triton College in River Grove, Illinois. Ten thousand square feet of space worked out just about right to display over 250 animals. Signage by Brian Jones's father added a professional tone to the event. In the background Brian's iguanas have staked out territories at the tops of tall branches and unconcernedly bask under heat lamps. A potential CHS member enjoys the show. Bob Bavirsha demonstrates how not to hold a snapping turde unless you're on very good terms with it. Lou Daddono's big Burmese pythons seemed content in flieir private swimmmg pool, despite flie lack of water. The pool was made available courtesy of Norbert Pools of Niles, Illinois. i 1 This hands-on tortoise display pen was designed and built by Brian Jones. 189

28 Bull. Chicago Herp. Soc. 29(8).190, 1994 Herpetology 1994 In this column the editorial staff presents short abstracts of herpetological articles we have found of interest. This is not an attempt to summarize all of the research papers being published; it is an attempt to increase the reader's awareness of what herpetologists have been doing and publishing. The editors assume full responsibility for any errors or misleading statements. JCM SNAKE MORTALITY ALONG ARIZONA HIGHWAYS P. C. Rosen and C. H. Lowe [1994, Biological Conservation 68: ] present an algebraic method for estimating highway mortality in snakes, based upon carefiil data collection during low-speed driving on paved roads. A total of 368 snakes (104 live, 264 dead) were recorded over four years on State Route 85 from Why to Lukeville, Pima County, Arizona, during 15,525 km of road cruising, mostly within Organ Pipe Cactus National Monument. The authors computed an estimate of 2383 snakes killed (13.5/km/year) during die four years on this stretch of pavement, and estimate that actual numbers killed would be closer to 4000 (22.5/km/year). Two taxa of special conservation interest, the Mexican rosy boa, Lichanura trivirgata, and the Organ Pipe shovelnose snake, Chionactis palarostris, appear to be relatively strongly impacted by highway mortality. Overall, along the 44.1 km transect, die estimated highway mortality amounts to the removal of 5 km' of snake population over the four years of smdy. It is clear that roadways, especially if paved, substantially damage snake populations. NESTING ECOLOGY OF THE BROWN CAIMAN IN CANO NEGRO, COSTA RICA J. Allsteadt [1994, J. Herpetology 28(1): 12-19] reports diat despite die widespread distribution of Caiman crocodilus fuscus {=chiapasius) in Central America, few detailed smdies of its reproduction are available. In northern Costa Rica, brown caimans nest in riverine forest adjacent to rivers or canals during the early wet season. Thirty-five nests were located and measured in ; detailed egg parameters were measured from 17 clutches. Mean incubation temperamre of nine nests averaged 31.8 C ( ''C). Individual nest temperamres ranged from 29.6 to 36.0 C. Egg temperatures increased by 3 C in five nests during the nesting season, but temperatures in four other nests remained nearly constant. Flooding accounts for high embryonic mortality by destroying entire nests; egg predation was minimal. Hatching occurs in September and October during high water levels when aquatic habitat is abundant. Delayed hatchmg of some nests suggest that a stimulus may be useful to initiate hatching. Typically, parent caimans opened nests to release the young. Hatchlings formed sibling groups near the nest site and often remained togedier for over 1.5 years. Mound nesting is favored by the optimal conditions of the wet season in Caiio Negro. Wet season nesting increases the availability of resources and suitable habitat to hatchling caimans. Abundant habitat during the wet season enables hatchlings to avoid predation and cannibalism. The predominant use of riverine forests for nesting by caimans in Cano Negro has important ecological consequences and management implications. BOTHROPS ASPER BITES AND FIELD WORKERS D. L. Hardy, Jr. [1994, Biotropica 26(2): ] smdies die problem of pit viper bites in field researchers working in Middle America (Mexico to Panama) using case histories of nine biologists and one local project employee bitten during the period of 1980 to All snakes involved were Bothrops asper. Based on local tissue and systemic effects, seven cases were severe, three with permanent disability. There were no fatalities. Although antivenin was administered intramuscularly as field treatment to seven victims and later intravenously as hospital treatment to nine as patients, its effect on outcome was uncertain. Nevertheless, intramuscular antivenin is recommended following an adult Bothrops asper bite when there are signs of envenomation and travel time to a treatment facility is more than four hours. The Extractor (Aspivenin) suction device may be of some benefit, but incisions, constricting bands and electroshock should not be used. Having a prior plan of action and implementing it effectively is most likely to improve outcome. Three bites occurred during 1.5 million person-hours in die field with Bothrops asper at four field operations in Belize, Costa Rica and Guatemala. Although risk of Bothrops asper bite to field researchers is low, it should not be ignored. THE BLINDSNAKE AND THE BULLDOG ANT J. K. Webb and R. Shine [1993, Animal Behavior, 45: ] report the Australian blindsnakes, Ramphotyphlops nigrescens, are small, fossorial snakes feeding on the larvae and pupae of ants. Examination of stomach contents show diat larger blindsnakes consumed larger prey items dian, and different prey species from, smaller blindsnakes. Only the largest snakes consumed brood of the large and formidable bulldog ants, Myrmecia. Three hypotheses for the causal basis of this size-related shift in dietary habits are tested. Observations on captive snakes suggest they are gape-limited predators, and the prey items eaten by small snakes in the field are as large as they could physically ingest in the laboratory. Hence, the absence of large prey items from small snakes may be due simply to gape limitation. However, medium-size blindsnakes are capable of ingesting items larger dian those diat ttiey take in die field. The absence of Myrmecia in die diets of these snakes is not due to prey choice (Myrmecia are readily eaten in the lab) or to an inability to locate or follow pheromonal trails of this ant genus, because all size classes of blindsnakes readily follow Myrmecia trails in the laboratory. Instead, the absence of Myrmecia from the diets of smaller snakes is probably due to die snakes' size-dependent vulnerability to nest defense by the bulldog ants. In laboratory encounters, Myrmecia workers are capable of biting and stinging (and evenmally killing) small blindsnakes, but larger snakes are relatively invulnerable to attack. 190

29 UnofHcial Minutes of the CHS Board Meeting, July 15, 1994 The meeting was called to order by President Marcia Rybak at 8:06 P.M. Board members Jim Caspar, Stacy Miller, Tony Rattin and Jack Schoenfelder were absent. Officers' Reports: Minutes of the May 13 board meeting were read and approved. Treasury: There is approximately $21,000 in die CHS treasury. Treasurer Gary Fogel announced the CHS can no longer operate as diough it has unlimited fimds. He made several suggestions for saving money including deferring the grants program and cutting back on the cost of monthly programs by using more local speakers. Mike Dloogatch suggested that die CHS has recently begun several initiatives to raise money, and that we can always shorten the length of the Bulletin by several pages each month to save money if necessary. Membership: There are currendy 1,854 CHS members. There was a discussion regarding possible reasons for nonrenewals. Ron Humbert suggested we send a postcard to nonrenewing members, asking the reason for the nonrenewal (e.g., dues increase, new location inconvenient, lack of regular book store insert) and advising fliem diat the book store is back in operation. Jill Horwich will draft the form for this postcard. Director of Sales Report: Joan Moore reported on die CHS Bookstore operations at the 1994 CHS Expo and at the June meeting. After encountering many problems with the new Booklog software, Joan Moore believes the CHS now has a stable, working version ready to go. Assuming all goes well, there will be a booklist in the August Bulletin, with regularly appearing booklists thereafter. In exchange for in effect using Joan to test die new version of the software, die company has agreed to make some custom modifications in our package free of charge. Committee Reports Taxes: The CHS federal income tax reports for 1993 have been completed and filed. Monflily Raffle: Marcia Rybak presented die results of die raffle held at die June meeting. Expo: Gary Fogel reported that the CHS netted approximately $1,500 at the 1994 Expo, excluding new memberships and merchandise sales. Ron Humbert will chair the Expo committee for the coming year. The 1995 Expo is scheduled for May 20-21, 1995, at Triton College. It will be relocated ft^om die basement to the fu-st floor gymnasium, which has better lightmg, no central pillars, higher ceiling, better temperatureregulating capability and a large cargo gate nearby. The room is approximately the same size as the basement room that housed the 1994 Expo. Anyone interested in participating in the 1995 Expo committee should contact Ron Humbert. Adoptions: Ben Entwisle reported that adoption activity has been high, with many people wanting to adopt animals. For those interested, there are plenty of iguanas in need of homes. Midwest Conference: Char Haguewood is checking with various hotels in the area as to availability to host the 1995 Midwest Herpetological Conference. The first committee meeting to plan the conference was scheduled for July 22 at Mike Dloogatch's downtown office. Members interested in helping to plan the Conference should contact Char Haguewood. Shows: The annual Chicago Academy of Sciences show is scheduled for September 24 nvi 25. The theme is "Our Weakening Web - The Story of Extinction." Oz Park Festival is scheduled for August 6-7. One person is needed each day to man die CAS table at this event. Mail order operations: The feasibility committee has been put on hold tmtil all the software problems have been solved and booklists are published on a regular basis. New Business: Show Policy: Marcia Rybak presented a draft of an official CHS show policy. She will compare this draft to the previous CHS show guidelines and come up with a show policy for action by the Board. Location: Holding general membership and board meetings at die Field Museum is costing die CHS in excess of $5000 annually. Marcia Rybak suggested forming a committee to explore die possibility of a new meeting location. The meeting adjourned at 9:45 P.M. Respectfully submitted, Jill Horwich, Recording Secretary SERPENT CITY, inc. S^fAKE HUSBANDRY SINCE I9SS DISPOSABLE REPTILE CAGE LINERS State-of-the-Art Cage Sanitation Huge labor savings over newspapers or wood shavings 100% biodegradable safe, non-toxic, no dyes FLEXWATT HEAT TAPE CAPTIVE-BRED SNAKES 30-day guarantee FROZEN FEEDER RODENTS HUSBANDRY EQUIPMENT CALL OR SEND S.A.S.E. FOR FREE1994 CATALOG SCOTT J. MICHAELS, DVM P.O. Box 657 Island Lake, IL USA*

30 Advertisements Accrue more herpetological knowledge: Connect with your peers throughout North America and abroad. The HERPETOLOGY ONLINE NETWORK is active 24 hours/day. Any computer modem can access Herp-Net via (215) ( baud 8-N-l) or (215) (9600-(- V32, V42bis.) Submit news via FAX: (215) any time. Attention: the Varanid Information exchange is a society of herpetoculturists sharing an interest in MONITOR LIZARDS. Members receive the bimondily newsletter, VaraNews. Annual membership is: U.S., $10; foreign, $12 surface/$15 air. For a free copy, send a legal-size SASE to: Varanix, 8726D S. Sepulveda Boulevard, #243, Los Angeles CA BOA SURVEY: Please write for my questionnaire on Boa constrictor reproduction. Even if your animals have not reproduced, please respond if they are at least four years old and have had the opportunity. In return for a completed survey you will receive a chart showing the subspecies, their scale counts and range. William Joy, P.O. Box , Arlington TX INTERNET: @COMPUSERVE.COM Contest: Why I love tordes and tortoises stories and/or drawings. Everyone is a winner. Winners receive free one-year membership and subscription to "Tyler's Tortoises, Terrapins and Turtles," P.O. Box 8181, Flint MI Please submit entries by December 31, All entries become property of TTTT and may be published in newsletters. For sale: WILLABD'S RODENT RANCH'em Ben sent ya! Live or fresh-frozen mice and rats-mice; pinks to hoppers $.35 each or 3/$1.00; weanlings to adults $.50 each; jumbos $.75 each rats; pinkies to chubbies $.50 each; fuzzies $1.00 each; small $1.50 each; medium $2.25 each; large $3.00 each; jumbo $4.00 each. Discounts available on large one-time orders. Prices subject to change without notice, availability may vary. Pick-up or delivery at the CHS meeting, shipping available for large frozen orders. Contact Mike Miller (days) at (708) For sale: fflgh QUALITY FEEDER ANIMALS PRODUCED FROM THE BEST LAB DIETS AVAILABLE. Tenfli year of production and supply of frozen feeder animals. All feeders can be removed one at a time from the bag; they are not frozen together. All orders will arrive frozen. Now offering seven sizes of mice: small newborn pinks, medium size pinks, large fiizzy pmks, extra large fiizzies/small hoppers, juvenile mice, young adults, and large adults. Also available are pinkie rats, baby chicks, and quail chicks. Orders sent special delivery, postage extra. Free pricelist. Kelly Haller, 4236 SE 2501 Street, Topeka KS 66605, (913) , after 6 P.M. Central Time on weekdays, all day Saturday and Sunday. For sale: THE GOURMET RODENT: rats and mice pinkies, fiizzies and adults. Quantity discounts. PleaSe send a SASE for pricelist or call BUI Brant, 6115 SW 137di Avenue, Archer FL 32618, (904) For sale: top quality mice, rats and Chinese dwarf hamsters. All sizes at competitive prices. Gary W. Allison, 919 Wyandotte Street, Bethlehem PA 18015, (215) For sale: murine-pathogen-free rats and mice available in all sizes, live or frozen; pinkies, fiizzies, crawlers, small, medium and large. Frozen crawler mice in lots of 2000, $.17 each. Also available, full grown hairless mice. FOB shipping point. Master Card accepted. Call (518) between 8:00 A.M. and 5:00 P.M. or write SAS Corporation, 273 Hover Avenue, Germantown NY for prices and additional information. For sale: high quality feeder mice. Shipped UPS Next Day Air. All mice are properly processed to insure a quality product. Fourth year of production and supply of frozen feeder mice. Prices; pinks, $25/100; fuzzies, $30/100; weanlmgs, $35/100. Also available are 4 oz.-i-rats, $100/100. Quantity discounts available. The Mouse Factory, P.O. Box 85, Alpine TX 79831, (915) , Ray Queen. For sale: rodents and reptiles, Dallas-Ft. Wordi Metroplex. J.R.'s Cowtown Critters, 1002 Harrison Avenue, Arlington TX 76011, (817) 460-RATS. For sale: live or frozen feeder mice and rats, all sizes available, from pinkies to adults. Quantity discounts. Lowe Labs, (708) 749-RATS. For sale; Pearlco ceramic heating elements, come in 60w, loow and 150w, $25 each or 2/$48 plus shipping; drip systems for reptile and amphibian enclosures, two styles, $10-12 plus shipping. Frozen rodents always available. Mice: pinks-weanlings, 3/$l; subadult, 50C; adults, 60C; large adults, 75C. Rats: small, 75-90C:; medium, $ ; large, $ ; jumbo, $ Call on quantity discounts [note: discotmt on orders picked up at CHS meetings]. Live rodents available at a higher price catalog finally fmished-send SASE. Scott J. Michaels, D.V.M, Serpent City, P.O. Box 657, Island Lake IL 60042, (815) For sale: mice and rats of all sizes. Also frozen toads. Call or write for price sheet. Keith Simpson, Rt. 3, Box 903, Knox IN 46534, (219) For sale: I'm back, Steve's Rodent Farm. Rats only, small and medium, live or frozen, quantity discounts. Shipping available. Call or write for price list. Steve Waldrop, c/o Steve's Rodent Farm, Rt. 7, Box 1154, HaleyvUle AL 35565, (205) For sale: FROZEN RODENTS. Mice pinks, $25/100; adults, $35/100; breeders, $55/100; rats, $75/100 and up, jumbo available occasionally. KJ's Rodents, R.R. 1, Box 266, Deo-oit Lakes MN 56501, (218) For sale: frozen rats, quarter-pound rats, $1 each. Steven Bostwick, 901 4th Sti-eet SW, Altoona la 50009, (515) For sale: feeders etc mice: pinkies, $.50; fiizzies, $.65; adults, $l-rats; pinkies, $.75; fiizzies, $1.25; weanling, $1.40; snull, $1.75; medium, $2.75; large. $3,75; jumbo, $4.75. Reel Pets, 4741 W. North Avenue, (312) For sale: 42 frozen pinky mice, $15. I expected hatchlings but they never arrived. For pickup in Lakeview or River North areas. Austin, (312) , leave message. For sale; SUPERWORMS, Zophobias mono, ultimate food for reptiles, birds, fish; $18.95/lb., plus $3 shipping. Keeps 8-10 months at room temperature, far less chitin for complete digestability. Send SASE for information to Joe Sutter, P.O. Box 59119, Birmingham AL 35259, (205) , 4 P.M. - 7 P.M. Cenffal Time. For sale: large selection of captive bom reptiles and amphibians, books and supplies. Send S.A.S. business-size envelope for complimentary price list or $5 for a 1 year subscription (bi-monflily), to Twin Cities Reptiles, 540 Winnetka Avenue North, Golden Valley MN 55427, (612) For sale: reptile cage disinfectant and deodorant. Kills many types of infectious disease detrimental to herps. $3.91 for 2 oz (makes 1 gallon). Wholesale and quantity prices available. Make checks payable to Bengal Pets, Inc., 1088 Chapman Road, Jesup GA 31545, (912) For sale: overstock sale. Phillips plastic boxes (aka Wilson; Wilhold; Stillwell). Shoe boxes, $2.25 each or $25/dozen; high sweater boxes, $6.15. Serpent City, (815) For sale: HERPETOLOGICAL DIRECTORY. A valuable information source contaming private and commercial breeders, foreign exporters, U.S. and foreign herp societies, wholesalers, and sources for rodents & other food items, supplies & equipment and publications. Send $15 to FAUNA, 2379 Maggio Circle Unit C, Lodi CA 95240, (209) MC/VISA For sale: Tired of getting bitten while collecting, feeding or maneuvering animals? Try my Whimey tongs 10% discount to herp society members 2', $65 each; 2Vi', $66 each; 3', $67 each; VA', $68 each; 4', $69 each, 5', $85 each; 6', $90 each. Will ship U.P.S. extra. Send money order or cashier's check to Don Lunsford, 5661 Walkerton Drive, Cincinnati OH

31 Advertisements (cont'd) For sale: set of five posters first published by Army Air Forces Tactical Center for easy identification and habits of venomous snakes of the world, $9 postpaid; Field Guide to the Snakes of South Vietnam by Simon Campden Main, $10 postpaid; Batrachia of North America by E. D. Cope, $16 postpaid; Laboratory Anatomy of the Iguana by J. C. Oldham and H. M. Smith, $17 postpaid; Time of the Turtle by J. Rudloe, $5 postpaid; Atlas of Amphibians and Reptiles of New Zealand by C. P. Pickard and D. R. Towns, $18 postpaid. Catalog of over 4000 herpetological titles sent free upon request. Herpetological Search Service & Exchange, 117 E. Santa Barbara Road, Lindenhurst NY 11757, (516) For sale: collection of 120 different outdated antivenom/snakebite kits from all over the world, $6(X). Also, collector's item snake catching box made and used by Carl Kauffeld. This is the box Carl used on his couectmg trips to South Carolina, 30"l x 20"h x 12"w, three compartments, top opens and two side compartments. Box was never cleaned out and still has (unsoiled) Staten Island Sunday Advance newspapers dated April 17, See book Snakes and Snake Hunting for picmre of box, $500. Steve, (516) [NY] For sale: Rattlesnake Hunting, a 60-minute video by a 30-year veteran collector. Don't miss this one! It's as close to being in the field as you can get. (Not a slick commercial production.) Send check or money order for $19.95 plus $3 postage & handling to: D. Wheeler, 2705 Sunset Trail, Riverwoods IL Allow 4-6 weeks for delivery. For sale; 125-gallon aquarium, seam leaker, versa hood with two fluorescent strip lights, wood oceanic stand, $250 or best offer. Michael Flaherty, D.V.M., Elk Grove IL, (708) For sale; African pygmy hedgehogs. Young males, $150; young females, $200; young pairs, $300; four male and four female adult proven breeders, $700/pair or $2500/breeding group. Hedgehogs can produce three to four litters a year with 3 8 babies each litter. I would consider trading any of the above for reptiles. Richard Evans, West Texas Reptiles, (806) or (806) For sale: c.b. giant day geckos, $30; Bibron's geckos, $20. Also, a pair of 3-year-old green basilisks, $300 o.b.o. Jim, (312) , leave message. For sale: one male and one female Nicaraguan banded geckos {Coleonyx mitratus), $90/pair; one male and one female Madagascar day geckos (Phelsuma madagascariensis boehmi), proven breeders, $135/pair; one male and one female lemon geckos (Gekko uukouskif), $30/pair. Also, c.b. frog-eyed geckos (Teratoscincus scincus), $50 each. All lizards in top shape and have exceptional colors. Philip A. Tremper, Jr., P.O. Box 264, Clintonville WI 54979, (715) For sale: baby leopard geckos (Eublepharis macularius), c.b. '94 from beautiful zoo-bom parents, eight left, $25 each; 10" baby savannah monitor, (Varanus exanthematicus), beautiful color with lots of gold/orange, perfect health, deparasitized, $65 (includes 50 frozen mice). For local pick-up only. Rob, (708) For sale; bearded dragons (Pogorto vrmcfpy), $75. Quantity discounts. Randy Gray, (817) eves. [TX] For sale: Chinese water dragons {Physignathus concincinus), c.b. '93, $40, c.b. '94, $15; bearded dragons {Pogona vitticeps), red morphs, c.b. '94, $90. Marcia Rybak, 736 N. Western Avenue, #315, Lake Forest IL 60045, (708) For sale: rhinoceros iguanas; Cuban iguanas; green iguanas; Geochelone sulcata; G. elephantopus (Florida sales only). D. J. or Sam, (305) [FL] For sale: mrtles. Two Juvenile Chinese threadheads (Ocadia), $25; male Chinese box turde {Cuora flavomarginata), $40; two 3-4" female Suwanee cooters, $25 [permit needed in Illinois]; 4" female hieroglyphic cooler, $15 [permit needed in Illinois]; one male and two female 4" Florida redbelly sliders, $45/lot; six 2-3" Barbour's map turtles, $40 each; two male spotted turtles, $45 each [permit needed in Illinois]; two male and two female Russian tortoises, flawless, $100/pair; one male and one female 5" ornate wood turtles {Rhinoclemmyspulcherrima manni), $100/pair; two hatchling ornate wood turtles, $75 each; 3" leopard tortoise, $125; 2" male Reeves' mitle, $15. Will trade for tortoises. Ed, (519) [Onurio, Canada] For sale; c.b. hatchlings. Ceylon pond turtles (Melanochelys trijuga thermalis), $50; Mexican milksnakes {annulata), $55; Honduran milksnakes, $90; eastern milksnakes, $10; Arizona mountain kingsnakes {Lampropeltis p. pyromelana), $100; Huachuca mountain kingsnakes (L. p. woodini), $100; graybanded kingsnakes, $95; Queretaro kingsnakes {ruthveni), $90; California kingsnakes, $20; Baird's rat snakes, $35; com snakes, $10. Henry Cohen, 24 St. Johns Place, Buffalo NY 14201, (716) For sale; one adult male Guatemalan milksnake {Lampropeltis triangulum abnorma), c.b. '92, $125. Jeff, (708) For sale: anerythristic Honduran milksnakes, $1000, heteros $500, possible heteros $ Large selection of locality-bred altema Loma Alta, Langtry, Juno Road, River Road, $75-150; Stuart's milksnakes, $125; Val Verde County annulata, $85; Sinaloan, Pueblan and Mexican milksnakes, $50-75; "The Geographic Variation of the Gray-banded Kingsnake," a color poster, $10 each plus $2 shipping. Dan Johnson, P.O. Box 42505, Houston TX , (713) For sale: repdle enthusiast located in mid-missouri offering a wide selection of captive-hatched snakes for sale this year. Corns, kings including Arizona and Mexican {knoblochi) mountain kings, Nuevo Leon, grey-banded, San Luis Potosi kings and more. Milksnakes such as Mexican, Pueblan, Honduran (including tangerines), nelsoni and arcifera; gopher snakes Pacific striped regular and albinos, albino Sonorans; western hognose [permit needed in Illinois]; regular, hetero and albino Burmese; boas and many more will be available. All breeder snakes were hand-picked as juveniles and owner has over 10 years of experience with many types of reptiles and flieir breeding requirements. Also available fliis year after 3 years of successful breeding axanthic trans-pecos rat snakes. These beautifiil grey/blue snakes were derived from wild-caught adults and are now well established. Also, two male Mexican mountain kings (Knoblochi), c.h. '91 & '92, $300 each; two central plains milksnakes (Lampropeltis triangulum gentilis), one female, one sex-unknown, captive raised from juveniles and gorgeous, $75 each. All animals are guaranteed healfliy and feeding on rodents. Unrelated animals available for many species and quantity pricing is always available. Send SASE or call Troy at 1954 State Road F, Fulton MO 65251, (314) For sale: female norfliem pine snake, c.b. '93, nice b/w contrast, $65. (708) For sale: one male and one female c.b 1991 albino San Diego gopher snakes (Pituophis melanoleucus annectens), proven breeders, $175/pair; one male and one female amelanistic com snakes, c.b. '91/'92, $100/pair. Jeff, (312) For sale: surplus animals. Two male and five female Mexican milksnakes; two male and four female albino California kings; two male and four female Pueblan milks; one male and four female Mexican black kings; two male and fliree female tangerine Honduran milks; two male and four female albino Sonoran gophers; one male hetero Sonoran gopher; one male and one female snow corns; one male and one female creamsicle corns; one male and one female albino corns; one male and one female normal corns; two male and two female San Luis Potosi kings; one male and one female albino prairie kings; one hetero female prairie king; one normal female prairie king; tiiree male and fliree female western hognose [permit required in Illinois]; one male and one female jungle carpet pythons. Most of flie above are adult animals and several are proven breeders. Prefer to trade for odier adult animals (leopard geckos, carpet pythons, Borneo blood pythons, and other colubrids and holds), but will consider cash offers. Richard Evans, West Texas Reptiles, (806) or (806) For sale: c.h. 5/15/94 ball pyflions, $40 each; c.h, 7/29/94 com snakes, red albinos, $20 each, snow corns, $25; one juvenile black and white banded Cal. king, $35. Also, 10-gallon aquarium widi hood and solid oak stand, like new, $40. Can deliver to CHS meetings. Art Nohlberg, (312)

32 Advertisements (cont'd) For sale: for Chicago area pickup only, c.b. '93 Kenyan sand boas, feeding on live ftizzy mice, $55; c.b. '93 banded California kingsnakes, mixed parents, feeding on dead chubby mice, $25. Janice, (708) , please leave a message. For sale: 5' female albino Burmese python with 40-gallon tank, $400 o.b.o. Ron, (708) For sale: large female Burmese python, very mellow, good eater, needs good home, $125 w/ cage o.b.o. Marcie, (217) [Champaign IL] For sale: diamond pythons, c.h. 6/24/94; Dumeril's boas, c.b. 5/18/94 and 6/25/94; western hognose hatchlings, all from unrelated parents [permit needed in Illinois]. Also Okeetee corns and leopard geckos. Eric Skov, HC 60 Box 175, Wells NV 89835, (702) For sale: 6' male D'Albert's python, 2'^-3 years old, $250. Bob Futo, (708) For sale: one male and one female green tree pythons (Morelia viridis), female 1.2 kg 5'A'±, virtually SOLID LIGHT SKY BLUE with a few scattered rusty spots, male 0.8 kg 5'±, dark lime green with a thin solid sky-blue vertebral stripe and several light spots, some yellow on snout, both feeding on rats and/or large chicks, c.b., c.r., super tame and gende, about ^'A-TA years old, $4350/pr; male green tree python {Morelia viridis), 0.5 kg 4'±, bright lime green with some scattered white and yellow spots, has retained about 10% yellow coloration, c.b., c.r., rat feeder, tame and ready to breed, $1500. Fred Wilson, (602) [Tucson AZ] For sale: BLOOD PYTHONS. Are you tired of trying to sort through the confusing and most likely erroneous geographical labels attached to the various color phases of Python curtusl You should be! Save yourself the mental anguish of trying to determine whether what you want is Sumatran, Malaysian, or Bomean in origin. If what you want are correcdy sexed and feeding blood pydion hatchlings from RED parents then you need to look no further. I have 3 clutches of blood eggs currently incubating. Hatchlings will be unrelated and from what many consider to be die finest Python curtus breeding stock in die land. Don't take my word for it, ask around. If you want to cut through all the "dealer hype" about the country of origin and color phase of die bloods you purchase, call me. You will receive great looking animals from red parents. No doubts about dieir origin. The best come from Roanoke, Virginia. Hatchlings are $400 each. Write to: David Lawrence, P.O. Box 6628, Roanoke VA 24017, or call (703) and leave a message if I'm not around. For sale: '94 babies. Carpet pythons, $150 each; Dumeril's boas, $350 each; rough-scaled sand boas, $ each; javelin sand boas, $100 each; Kenyan sand boas, $100 each. For free price list call or write Diablo Valley Reptile, P.O. Box 23246, Pleasant Hill CA 94523, (510) , % Chris Reimann. For sale: Bimini boas, Epicrates striatus foster, c.b. 9/93, $250/pair. Well started feeding on pinkies, will consider trades. Jim, (305) For sale: genetic striped boas? Normal adults produced 6 normal, 7 partial striped and 6 fully striped babies on 6/12/94. These are not heat mutations. Prices range from $300 to $800. Steve, (206) [Seattle] For sale: Guyana redtails, c.b. 4/26/94, eating well, nice colors coming in, $500/pair; NW Peruvian boas, c.b. '94, males only, blue/gray heads, bold black head spear and black tail blotches, $300 each; Borneo blood pythons, unrelated available from 5 clutches, eat well, grow fast, nice docile animals, $ each; black blood pythons due early Sept., our second year, most of last year's are very black, $350 each; two male black blood pydions, c.b. 10/1/93, $350 each; two adult female brown water pythons, $275 each or both for $500; one male and one female yeariing ball pythons, eat well, $100/pair; male ball pyflion, c.b. '90, nice, $75. Tim or Janice, (903) [TX] For sale: beautiful, bouncing baby Brazilian rainbow boas, three males and six females, excellent specimens, bom 7/10/94, mahogany parent female and brilliant orange parent male, both have high iridescence, all babies vet checked and feeding, $225 each. Garry Roberts, (708) For sale: moving and must sell 6' male redtail boa, $100; breeding pair of ball pythons, 4' male, 5' female, $140/pair; 2' albino com snake, $40. All in excellent health. (608) [Madison WI] For sale: three male and four female c.b. '94 red Lamar-strain Brazilian rainbow boas, outstanding specimens, $350 o.b.o.; two male and two female c.b. '94 jungle carpet pythons, $400 o.b.o.; pair of c.b. '93 diamond x jungle carpet pythons, $800 o.b.o. (703) , please leave message. For sale: two male boa constrictors, c.b. 7/91 and 6/92, $70 each. Jack Phillips, (708) For sale: Between July and September I expect young from the following boid species sand boas, Eryx colubrinus loveridgei, $60; E. conicus, $75; E. j johni, $150 rosy boas, Lichanura trivirgata saslowi, $150; L. t. trivirgata, $75 Dumeril's boas, $300; olive pydions, Morelia o. olivaceous, $1500. Also, surplus are one male and one female adult European sand boas, Eryx jaculus familiaris, $275/pair; one male and two female adult Tatar sand boas, E. tataricus vittatus, $150; young male ('93) Dumeril's boas, $300; one male and one female 9-10' amethystine pyuions {kinghomi), $1000/pair; one male and one female 7-9' Bameck amediysrine pydions, $700; 7' male Moluccan amethystine pydion, $175. David Sorensen, 4100 N. 78th Street, Milwaukee WI 5322, (414) For sale: baby Argentine boas (Boa constrictor occidentalis), bom 7/28/94 from nice, unrelated parents, $ each. Rob Nadian, (216) , leave message. [OH] For sale: Argentine boa constrictors (Boa constrictor occidentalis), c.b. 8/94, parents are '86 imports, unrelated to all otiier U.S. bloodlines, $750/pr; Surinam red-tailed boas, '94 babies, above average color, $ each; 3' male Surinam redtail, bom here 9/93, exceptional color, $500; Guyana redtail boas, from exceptional adults, due mid-september, inquire; Brazilian rainbow boas (Epicrates c. cenchria), due mid-september, $250 each; spotted pydions (Liasis maculosus), hatched 6/94, well-started, $ each; albino Burms call for price; many colubrids too numerous to listmountain kings (zonatas, pyros) and milks (triangulum) in particular. Call or send business-size SASE for updated summer snake/supply catalog. Serpent City, P.O. Box 657, Island Lake IL 60042, (815) For sale: Macklot's pythons, Liasis mackloti, c.b. '94, $125 each; Amazon tree boa, c.b. '93, $125; one male and one female Kenyan sand boas, Eryx colubrinus loveridgei, c.b. '93, $150/pair; one male and one female Burmese pythons, 8', c.b. '91, $200/pair; Peruvian red-tail boa. Boa constrictor ssp., c.b. '92, locality data on parents, $300; male albino Burmese pydion, c.b. '93, hetero for albino-green, $200. Call for free price list. Steven Bostwick, 901 4th Street SW, Altoona la 50009, (515) For sale or trade: one adult male Paroedura pictus, long term captive, striped phase, proven breeder, $60. Brett DePoister, R.D. 2, Box 2158, Fleetwood PA 19522, (610) Free!!! NATION'S LARGEST CATALOG of reptile and amphibian supplies at the guaranteed lowest prices. See our display ad in diis Bulletinl Big Apple Herpetological's 48-page catalog includes incubators, cages, heating, digital thermostats & diemometers, lighting, cage accessories, husbandry & feeding/dosing equipment, medications, vitamins, foods, herp shirts and books. We also sell a large variety of reptiles. For FREE catalogs and product orders: (800) For product information: (212) For reptile information and sales: (212) Visa, MasterCard, Discover accepted. BIG APPLE HERPETOLOGICAL, P.O. Box 632, New York NY

33 Advertisements (cont'd) Pet-sitting service: Complete in-home pet and plant care, specializing in reptiles and other unique pets. Dogs and cats welcome with open arms, midday dog walking. Experienced and dependable service by devoted animal lovers, serving Chicago and suburbs, reasonable prices, fully insured and bonded. Ask about our habitat consultation and custom-made cages. Exotics & More, (312) or page us at (708) Wanted: information on custom-built herp cages. I am looking for someone to create a perfect home for my two green iguanas who will soon outgrow their current home. Susan, (312) Wanted: businesses or persons interested in brokering for moderate-sized reptile breeder. See ad above for types available. Troy, (314) Wanted: male Travencore tortoise (Geochelone travencorica) for breeding purposes; Travencore information welcomed as well. Leave a message on the "ARK line," (604) or write: ARK - Association of Reptile Keepers of British Columbia, Tarrah Satchwell, Sapperton RPO, Box 43020, New Westminster, British Columbia V3L 5P7, Canada. Wanted: snake-necked turdes (Chelodina spp. and Hydromedusa spp.). Fred Caporaso, Natural Sciences Division, Chapman University, Orange CA 92666, (714) telephone, or (714) fax. Wanted: crocodile lizards (Shinisaurus crocodilurus); any female Indian star tortoises; juvenile "cherryhead" red-footed tortoises. John Park, (203) Wanted: adult male northwest Peruvian boa (black tail), also known as Tumbes boa. Tim, (903) [TX] Wanted: sand boas (Eryx elegans, E.j. jaculus, E.jayakari, E. Johnipersicus, E. muelleri and E. somalicus). David Sorensen, 4100 N. 78tit Street, Milwaukee WI 5322, (414) Line ads in this pubiieation are run free for CHS members $2 per line for nonmembers. Any ad may be refused at the discretion of the Editor. Submit ads to: Michael Dloogatch, 6048 N. Lawndale Avenue, Chicago IL 60659, (312) evening telephone or (312) fax. INCUBATORS DON'T MISS THIS! Get the Nation's Largest Catalog of Reptile & Amphibian Supplies Guaranteed Lowest Prices! > FREE 48 Page Catalog ^ THERIVlOSTATS Big Apple Herpetological (Formerly Big Apple Snakes) For Catalogs & Orders (800) For Product Information (212) Big Apple HerpetoIogical-P.O. Box 632-New York, NY I WE WELCOME RETAILERS! ASK FOR OUR WHOLESALE PRICE LIST! I RATS AND MICE Bill & Marcia Brant 6115 SW 137th Avenue Archer, FL (904) FAX (904) ISAII Rights Raaval HUSBANDRY MEISiCATIONS Full line of herpetological supplies Call for catalog 195

34 News and Announcements DONATIONS TO THE JUNE 29 RAFFLE The following is a listing of those individuals and businesses who generously donated items for our monthly raffle at the June 29 meeting. The donated items are shown in parentheses. Aquascape, Inc. Bannockbum IL (Critter Cage); Jill Horwich (turtle T-shirts); Wardley (Reptile Ten food); Ghann's Cricket Farm (cricket gift cenificate, high-calcium cricket diet); Reptile News Press {Basic Reptile Care, Safe Soap cage cleaner, monitor mug); Critter Company (herp bumper sticker); Terrarium Art (large snake cave); Coral Reef Pet Center (dog food); Tetra Terrafauna (snake poster); Tony Janowski (reptile cage carpet); Serpent City-Scott Michaels (plastic shoebox, cage liners); California Zoological Supply (canned herp food); Tom Zaremba (small tank with locking top); Ilene Sievert (plants); Fluker Farms (freeze-dried crickets); Valentine Equipment (heat pad); ZooMed (Habba Tree); Mardel Labs (Derma Shell); Grubco (mealworm gift certificate); Rep-Cal (calcium supplement); Advanced Vivarium Systems {The General Care and Maintenance of Bearded Dragons); Ron Humbert {National Geographic 1986 mrtle issue); anonymous (thermometer). LOUISVILLE ZOO CHARTER BUS TRIP Register now for the Chicago Herpetological Society's Louisville Zoo charter bus trip. The trip is scheduled for October 23 and will feamre a behind-the-scenes tour of the zoo's state-of-the-art HerpAquarium. This relatively new building (approximately five years old) is divided into biomes, or ecological communities, each stressing the interrelationships of animals and plants. The modern technology within the facility re-creates the climates as well as the sights and sounds of desert, forest and aquatic biomes. For example, the rainforest exhibit comes complete with simulated rain showers and hot, sultry temperamres. The motorcoach (equipped with reclining seats, six television monitors, VCR, stereo and rest room) will depart the Field Museum of Namral History's north parking lot at 5:00 A.M. and will retorn between 10:00 and 10:30 P.M. Registration is limited to the first 47 paying individuals. Tickets are $35 each (nonrefundable) and will be on sale at the general meetings. If you wish to mail your payment, please send a check or money order, payable to the CHS, to Anthony Rattin, Avon Court, Wildwood IL Be sure to include a current return address so that tickets and detailed information brochures can be forwarded to you UPSTATE HERPETOLOGICAL ASSOCIATION GRANT The Upstate Herpetological Association (UHA) is now accepting applications for a grant in the amount of $500. This grant is intended to support herpetological conservation, education or research. All proposals must be received on or before September 15, The successful applicant will be notified on or before November 15, Applicants need not be a member of UHA nor a professional herpetologist but shall be able to demonstrate a sincere interest in herpetology and the ability to complete the endeavors for which the grant is awarded. Successful applicants will be expected to submit a report of their work for publication in the association's newsletter, "Herp Beat" and/or give a presentation to either the Syracuse or Albany chapter of the UHA. It is expected that the successful applicant will acknowledge the UHA's support in any publications that result in part or wholly based on funding by UHA. Applications should include the purposes for which the grant is requested, the uses to which the grant will be put, and enough background information to enable reviewers to assess the applicant's ability to accomplish the objective for which the grant is requested. All applications should include a budget for the project and a completion date for the project. Applications should be held to a maximum of five pages, typed, double spaced. The UHA reserves the right to reduce the grant, divide the grant between two or more individuals or make the grant to an organization as a whole. The reviewers will be instructed to consider all proposals but may exercise a bias toward original research. New York herpetology, and captive husbandry and propagation of reptiles. Applications should be submitted to UHA, % Ed Schoene, D.V.M., HCR 68, Box 30B, Springfield Center NY Questions should be directed to Dr. Schoene, (607) CENTRAL ILLINOIS HERPETOLOGICAL SOCIETY ANNUAL SHOW On August the Central Illinois Herpetological Society will hold their annual herp show in the Lakeview Museum in Lakeview Park in Peoria, Illinois. There will be no selling of animals at this show. Hours will be 10-4 Samrday and 12-4 Sunday. For more information or for directions call Don Ay res, (708)