Wi A R ^BULLETIN OF THE US |S$N. 0025-4231 TRarylano f)tcpetologtcaI ©octety Department of Herpetology The Natural History Society of Maryland, Inc, MdHS . A FOUNDER MEMBER OF THE Eastern Seaboard Herpetological League DECEMBER 1984 VOLUME 20 NUMBER 4 Bulletin of the Maryland Herpetological Society Volume 20 Number 4 December 1984 CONTENTS ON SOME ASPECTS OF THE HISTOCHEMISTRY OF THE ALIMENTARY CANAL OF THE TERRAPIN, Mauremys caspica (GMELIN) (REPTILIA, Testudines , Emydidae) . Noo ry T . Ta i b 123 EFFECTS OF RATTLESNAKE ( Crotalus viridis oreganus ) ENVENOMAT ION UPON MOBILITY OF MALE WILD AND LABORATORY MICE ( Mus musculus) . William K. Hayes and Joseph G. Galusha 135 EFFECT OF MALE COURTSHIP ON STR I KE- I NDUCED CHEMOSENSORY SEARCHING IN A FEMALE URACOAN RATTLESNAKE ( Cvotalus vegrandis) AT NATIONAL ZOO . David Chiszar, Joan Chiszar, Trooper Walsh, Bela Demeter and Hobart M. Smith 1 4 5 NOTES ON THE ENIGMATIC Barisia imbricata OF THE BRITISH MUSEUM, AND ON ITS COLLECTION OF REPTILES FROM AMULA, GUERRO, MEXICO . Hobart M. Smith 152 REPTILIAN PARTHENOGENESIS . Cliff H. Summers 159 A NEW SUBSPECIES OF Centrolenella orientalis (ANURA: CENTROLEN I DAE) FROM TOBAGO, WEST INDIES . Jerry D. Hardy, Jr. 1 65 NEWS AND NOTES: THE HERPETOLOGICAL WORKS OF JACOB KLEIN. WITH EMPHASIS ON THEIR PERTINENCE TO MEXICAN HERPETOLOGY . John Johnson, Malvin L. Skaroff, Hobart M. Smith and Rozella B. Smith 174 SMITHOS I AN INSTITUTION - OFFICE OF FELLOWSHIPS AND GRANTS ANNOUNCEMENT . 190 NEW BOOK NEWS: PHYLLIS, PHALLUS, GENGHIS COHEN, S OTHER CREATURES I HAVE KNOWN . 191 THE TURTLES OF VENEZUELA . 192 HERPETOLOGY OF ARABIA . 193 NEWS AND NOTES: LITERATURE: FAUNA CLASSIFIEDS . 194 The Maryland Herpetological Society Department of Herpetology Hatural History Society of Maryland . Inc. 2643 North Charles Street Baltimore. Maryland 21218 BULLETIN OF THE Volume 20 Number 4 December 1984 The Maryland Herpetolog i cal Society Department of Herpetology, Natural History Society of Maryland, Inc. Bulletin Staff Executive Editor Herbert S. Harris, Jr. Steering Committee Donald Boyer Herbert S. Harris, Jr. Frank Groves Jerry D. Hardy, Jr. Jeff Thomas Officers President . V i ce-Pres i dent Secretary . Treasurer . Richard Czarnowsky Beth E . Cline Les Chaikin Les Chaikin Library of Congress Catalog Card Number: 76-93458 Membership Rates Full membership in the Maryland Herpetol ogical Society is $12.50 per year, subscribing membership (outside a 20 mile radius of Baltimore City) $10.00 /year, Foreign $12.00/yr. Make all checks payable to the Natural History Society of Maryland, Inc. Meet ings The third Wednesday of each month, 8:15 p.m. at the Natural History Society of Maryland (except May-August, third Saturday of each month, 8:00 a.m.). The Department of Herpetology meets informally on all other Wednesday evenings at the NHSM at 8:00 p.m. ’ ■ / ' Bulletin of the Maryland Herpetological Society Volume 20 31 December 1984 Number 4 On Some Aspects of the Histochemistry of the Alimentary Canal of the Terrapin, Mauremy s caspica (Gmelin) (Reptilia, Testudines , Emydidae) Noo ry T . Ta i b Abstract The distribution and localization of carbohydrates, lipids, nucleic acids, proteins and nine digestive enzymes in the epithelium of the alimentary canal of Mauremye caspica were studied. The histochemical reactions revealed the occurrence with variation of numerous active secretions of mucoscubstances and digestive enzymes. These findings are discussed in the context of its feeding habits. I ntroduct ion Mauremys caspica (Gmelin), the strip-necked terrapin inhabits both fresh and brackish water and occurs in northeast Saudi Arabia and several other countries of the Arabian Gulf (Loveridge, 1955). A survey of the literature revealed that little histochemical work has so far been carried out on the alimentary canal of reptiles (Wright et al., 1957; Anwar and Mahmoud, 1975; Chou, 1977; Taib, 1981; Suganuma et al., 1981; Taib and Jarrar, 1983) and no histochemical work had been undertaken on the alimentary canal of Mauremys caspica. The morphology and histology of the alimentary canal of Mauremys caspica were described in an earlier paper (Taib and Jarrar, 1982). The present study is an attempt at understanding the digestive physiology of this species in relation to the epithelium secretions of its alimentary canal . Materials and Methods Ten adult Mauremys caspica were used in the present study. Each animal was anaesthetized with chloroform and the following parts of the alimentary canal were identified, flushed of and isolated, namely, upper oesophagus, lower oesophagus, stomach (cardia and pylorus), small Bulletin Maryland Herpetological Society Page 123 Volume 20 Number 4 December 1984 intestine, colon, and rectum. Paraffin as well as unfixed fresh cryostat sections (5-12 ]im at -25°C) were routinely used. A tabulated outline of the h i stochemi cal methods employed is given in Table I. The control in each case consisted of parallel incubation of sections in media lacking a specific substance or using heat- i ncubat i ng sections. Visual estima¬ tion of the dye deposited in different regions using light microscopy examination was used as a measure of the relative activity of the enzymes and presence of other compounds (Figures 1-12). Table I List of H i stochemi cal Tests Undertaken Test or Technique Reference 1. Best 1 s carmine Best (1906) 2. Periodic acid-Schiff (PAS) Gurr (1962) 3. Diastase treatment (one hour at 37°C, Subsequently stained with PAS) McManus and Mowry (1964) 4. A1 c ian blue pH 2.5 Mowry (1956) 5. A1 c ian blue pH 1.0 Mowry (1956) 6. Alcian blue pH 2.5/PAS Mowry and Winkler (1956) 7. Alcian blue pH 1.0/PAS Mowry and Winkler (1956) 8. Osmium tetroxide Mallory (1961) 9. Sudan Black B Chiffelle S Putt (1951) 10. N i nhydr i n- sch iff Yasuma and Ichikama (1953) 11. Mercuric Bromphenol blue Pearse (1972) 12. Tol u i d i ne blue Pearse (1972) 13. Acid phosphatase Pearse (1972) 14. Alkaline phosphatase Gomori (1952) 15. Lipase Gomori (1952) 16. Nonspecific esterases Gomori (1952) 17. Exopept i dase McCobe and Chayen (1965) 18. Endopept i dase Yamada and Ofugi (1968) 19. Beta-gl ucuron i dase Hyashi et al . (1964) 20. Carbonic anhydrase Hausler (1958) Page 124 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December 1984 Figure 1 - Stomach. Periodic acid-schiff, X90. Figure 2 - Colon. Alcianblue pH 2.5, X90. Figure 3 Oesophagus. Toluidine blue, X120. Figure A - Stomach. Mercuric bromophenol blue, X90. Figure 5 - Stomach. Osmium tetroxide method for lipids, XI 20. Figure 6 - Stomach. Sudan Black B, X90. Figure 7 - Stomach. Lead nitrate method for acid phosphatase, X90. Figure 8 - Duodenum. Calcium-cobalt method for alka¬ line phosphatase, X90. Bulletin Maryland Herpetol og i cal Society Page 125 Volume 20 Number 4 December 1984 Figure 9 - Stomach. The a-naphthol acetate method for fionspeciflc esterase, X90. Figure 10 - Duodenum. McCobe and Chayen method for am I nopeptidase, X90. Figure 11 - Stomach. Ha'usler method for carbonic anhydrase, X90. Figure 12 - Stomach. Naphthol AS-BI Glucuronide method for 6-glucuronidase, X120. RESULTS The histological structure of the al imentary canal of Mauremys caspica has been described by Taib and Jarrar (1982). Accordingly, the oesophagus is lined with ciliated columnar and goblet cells and is without oesophageal glands. The surface epithelium of the stomach is of a single layer of columnar mucous cells with occasional goblet cells. The lining of the gastric glands consists of epithelial cells interspersed by mucous cells. The lining of the intestine is made up of both striated columnar and goblet cells. Throughout the intestines, goblet cells pre¬ dominate posteriorly. The result of these h i stochemi cal experiments carried out in this study are given in Table II. Ca rbohydrates : The lining epithelium of the oesophagus reacted strongly with carbohydrate-staining sequences. The results indicated the presence of neutral polysaccharides as well as sialomucins and sulfomucins. The lower oesophagus showed a prominent of sulfomucins while in the upper oesophagus, the amount of sulfomucins and sialomucins were almost equal. The apical surface of the ciliated columnar cells also stained for sialomucins. The covering epithelium of the stomach displayed abundant polysaccharides and little sialomucins and sulfomucins. Gastric glands were stained brightly with PAS. On treatment with diastase, a certain amount of PAS-positive material was removed from the gastric glands but the stomach lining remained unaffected. Page 126 Bulletin Maryland Herpetol og i cal Society TABLE II Results of Histochemical Reactions on the Lining Epithelium of the Alimentary Canal of Mauremys caspioa. - Volume 20 Number 4 December 1984 ON + I ! + + +1 +, +, + +1 ' 1 +» 1 1 1 + + + +1+1 +1 00 + + I + + I + • t t| + Jl tl + +l ' +" Jl ' + , +, + + + +I+II+H+I + I + 1+1 + + || I +11 +11 + +1 + I vO + i , + + + + + +i+i+i + +«+ + + + + + + +1 +1 +1 + + + + + +. + + +»• + +1+ + + I+ + + +I+ + + co cm + + + + + + + + + + | +| C- I + | c— + + + I I + + + + + + + 11 o* + + + + + + + + + + + + + | + n +| +| + +1 +u +1 +1 + + * ■K + + + + + + + + + + | +11 I +11 +| I +11 I +H +« P 3 <4H 4H •H X CJ CO I 3 •H CJ CO 3 O •H 4-1 0) PH ,p m O P o • • P *H CM H 4-1 JC 4-4 4H P. C 0 P 35 pa •H o p P P P- (X X s 3 PQ 00 •H a o rH •H a P p c n 4-i X 4*5 *3 u 3 3 \ PQ CJ p rH rH P P P P o X X *H CJ P rH CO 4-t •H •H PQ P CO P P •3 4-4 *3 4-4 a. P P 5>n 3 P CO 4-4 •H •H X CJ 3 P P CO a CJ P 4-4 rH 3 •H P rH rH •H p o 3 Q PQ < < 53 s H CP a) 3 •H X o u 4-» P % g CO o P CO P 4-> CCJ s: (X CO o -p cx 3 •H O < p co P 4-4 co X a co o X (X P p •rt rH CO X CO 0) CO co M P 4-4 CO CD >4-1 •H O 04 O- co P o 53 CO 0) CO co p- 0) CO CO 3 *H 4-1 p. 04 P. o p •H 04 CO CO 3 P. 04 a o 3 & 04 CO co u 3 X 5o P cO I CJ *H P o ■e cO CJ 04 CO CO 3 •H P O 4-i 3 O 3 t— l 00 I p 4-4 0) PQ CO u o •H 4-i 04 vO CO -3 P CO iH 00 CJ •H U 4-> CO CO O m co 3 U • o a r— I 3 50 4-4 P. CJ 04 • C«- <+ P 4-4 4-i P O P 3 c— 00 P o 4-i 4-4 CO V4 04 > + + + 00 p o 4-1 4-4 CO + + 04 4-4 P 4-4 04 • 3 «•> ON O P •H s *3 P * 4-4 O P rH + U O • 0k CJ X • P co • P 00 • n £ CO 0k 3 P 00 P + 1 P *rH • 0k X 4-4 X P. CO P O P P CO 4-4 p p ? O *rl 5h 4-4 4-t rH P p 1— 1 5 p 5> o a 0k X CO + 1! • 4-i • 0k CM O 4-1 •H P • »> 4-i P CO P CO 3 4-4 -O 00 CO P O p X PH A Pi o • 1 co r~~ • • P P O • O p •H 4-4 P 4-4 P *H O P. 4-4 P P. CO P 33 P 4-> 4-1 • P 4-4 r— 1 *H O 5o 04 5«5 * 04 3 o cj ■K * Bulletin Maryland Herpetolog i cal Society Page 127 Volume 20 Number 4 December 1984 In the intestine, the striated border of columnar cells and their basement membrane stained positively for neutral polysaccharides and sulfomucins. Goblet cells throughout the intestine contained neutral and acidic polysaccharides. However, unlike the goblet cells in the small intestine, those in the large intestine showed more intensive stain for sulfomucins. A predominant pattern of sialomucins in the goblet cells of the crypts of the intestine epithalium was also observed. Proteins: The lining epithelium of all regions of the alimentary canal gave positive reactions for proteins. The apical cytoplasm of the ciliated columnar cells of the oesophagus demonstrated the most pronounced staining. Gastric glands reacted strongly indicating the presence of a considerable amount of stored or enzymatic proteins. Globules of goblet cells showed faint 3~metachromas ia with toluidine blue, while their cytoplosmic granules gave strong $-metachromas i a reaction, indicating that the proteins were either glyco- or muco-protei ns . The nuclei of all ceils lining the alimentary canal were positively stained for proteins. Lipids: Most of the lipids were found in the cells lining the stomach, the gastric glands and the large intestine. Lesser quantities were detected in the oesophagus and in the large intestine. In the epithelial cells, lipid globules appeared to be concentrated in the basal region of the cytoplasm while their basal border as well as cells coat were densely packed with heavily staining globules. Goblet cells throughout the entire alimentary canal displayed a negative reaction for lipids. Phosphatases: A pronounced but diffuse cytoplasmic reaction of acid phosphatase was obtained in most of the cells lining the gastric glands. On the other hand, the cells lining the stomach showed only weak activity while a slighter reaction of acid phosphatase activity was noticed in the striated borders of the columnar cells lining the small intestine. Here, the reaction occasionally extended well into the distal half of the cell. Alkaline phosphatase demonstrated a moderately positive reaction in the gastric glands and the columnar cells lining the small intestine, but only slight activity of this enzyme was observed in the lining of the lower oesophagus and the anterior portion of the large intestine. However, the activity of alkaline phosphatase was less than that of acid phosphatase. Phosphatases are enzymes of lysosomal origin (Bowen, 1968) and are found in high concentration in the cells which are active in secretion (Reid, 1966). It is recognized that phosphatases are involved in supplying the energy required for the active process of secretion (Taib, 1976), and the Page 123 Bulletin Maryland Herpetolog i cal Society Volume 20 Number 4 December 1984 absorption in the duodenum in various rodents (Hugon and Borgers, 1968). The activity of alkaline phosphatase in the lining of the small intestine indicates a role in absorption of material from the lumen of the gut. Lipolytic activity: Nonspecific esterases activity was detected in the lining of all regions of the alimentary canal. The most pronounced reaction was also observed in the gastric glands. A pronounced reaction was also observed in the lining epithelium of the intestines with a gradual diminution of activity posteriorly. A low level of activity was recorded in the oesophageal lining. The goblet cells as well as the mucous-secreting portion of the cells lining the stomach showed no evidence of any esterase activity. Esterase activity has been demonstrated in both lysosomes and microsomes by biochemical and cytochemical techniques (Holt, 1963; Tappel , 1969). Gomori's Tween method failed to reveal any lipase activity any¬ where in the alimentary canal, although the validity of the method was successfully tested with sections of the pancreas. The failure to locate any lipase activity is probably due to its weakness or complete absence. It is probable that the unsaturated substrate (Tween 80) is attacked only by pancreatic lipase and not by other esterases. However, the Tween method for demonstration of lipases has not generally proved to be sufficiently sensitive unless the animals are maintained on a high fat-content diet (Jennings, 1962). Proteolytic activity: The lining epithelium of the small intestine gave an extremely strong and vivid reaction for ami nopept i dase with gradual diminution posteriorly. Considerable activity was also recorded in the gastric glands. The lining of the lower oesophagus, however, showed a faint positive reaction. The activity was almost absent in the large intes¬ tinal epi thel ium. Endopept i dase activity was moderately positive in the lining epithelium of the small intestine while a low activity was observed in the lining of the oesophagus, stomach, and gastric glands. In the large intestine the activity bacame noticeably almost negligible. It may be that these enzymes are probably concerned with the later stages of digestion and absorption of material from the gut lumen (Taib, 1976). Proteolytic enzymes have been recognized to be always present in the cells lining the gut. These enzymes when intracellular, are normally in an active state, often requiring partial digestion for activation (Smith, I960). Bulletin Maryland Herpetol og i ca 1 Society Page 129 Volume 20 Number 4 December 1984 Carbonic anhydrase: Intensive activity of carbonic anhydrase appeared in some cells lining the gastric glands. Considerable activity was observed in the lining of the lower oesophagus and the anterior part of the small intes¬ tine, but was found to be weak in the lining of the large intestine. Carbonic anhydrase is so far known to occur in an active condition and that it was associated with production of hydrochloric acid in the stomach (Jennings, 1962; Taib, 1981). This may perhaps explain the appearance of the intense reaction in the gastric glands which are known to be the source of gastric hydrochloric acid. 3-glucuronidase: Strong activity of 3~g1ucuronidase was indicated in the lining epithelium of the small intestine and it was localized in the distal parts of the cells. Some cells lining the gastric glands demonstrated moderate activity. A low level of 3-gl ucuron i dase activity was detected in the apical surface of the ciliated columnar cells of the lower oesophagus. The activity in the lining of the large intestine was almost negligible. 3~gl ucuron i dase is important in hydrolysing oligosaccharides and is known to occur in lysosomes (Novikoff, 1961), and in other parts of certain cells (Rosenbaum and Ditzion, 1963). D i scuss ion The result of this study showed that neutral mucosubstances are evident in the stomach while acidic mucosubstances occur in the oesophagus and the intestine. Sialomucins were most prominent in the lower oesophagus and sulfomucins in the large intestine. I may, therefore, postulate that acid mucosubstances may play some protective role against gastric reflux and aid in feacal discharge. The abundance of acid mucosubstances in the oesophagus of this species is, however, little understood since this terrapin usually feeds under water and it may take a large quantity of water to facilitate swallowing and no need of muco¬ substances secretion to be involved in this function. This pattern of mucosubstances distribution in the alimentary canal resembles that which occurs in several other reptiles such as the skink, Eumeces latiscutatus , the turtle, Glemmys japonioay the gecko, Gehyra multilata, and the snake Elaphe climaaophora (Chou, 1977; Suganuma et al., 1981). Epithelial mucosubstances occur both as intracellular components as well as extra cellular luminal secretions. Mucosubstances are believed to occur as glycoproteins when secreted into the gastrointestinal lumen (Kent, 1971). The diet of Mauremys caspica is varied (Mahmoud and Klicka, 1979) > containing plant material, insects, small invertebrates and amphipods. The feeding habits of this terrapin vary with the age of the animal and the seasons. The change in the dietary requirements with age being Page 130 Bulletin Maryland Herpetological Society Volume 20 Number 4 December 1984 partially related to their physiological demands. Juvenile terrapins have been observed to feed on small insects with a high calcium content which is necessary for shell growth (Mahmoud and Klicka, 1979). This terrapin feeds mainly on insects during early summer and with the decline in the insect population in late summer, it feeds more on plants. The result of this study shows that Mauremys oaspioa possesses an enzyme complement capable of breaking down a varied diet. The results also suggest that the considerable amounts of various enzymes in its stomach and the small intestine help to ensure the maximum utilization of the food ingested. Acknowl edgements I wish to express my appreciation to Mr. B. Jarrar for his assistance during the course of this study. 1 am indebted to Dr. P.H.D.H. De Silva for his helpful criticisms in the preparation of the manuscript. 1 am very grateful to the staff of the Photography unit in the College of Science for their assistance with the photographic work. 1 wish to thank Mr. M. Issa for technical assistance and to Mr. Z. E. Osman for typing the manuscript. L i terature C i ted Anwar, I.M. and A.B. Mahmoud 1975* "Histological and h i stochemi cal studies on the intestine of two egyptian lizards". Bui. Fac. So. Assiut. JJniv ., 4(1) : 1 01 - 1 08 . Best, F. 1906. "Uber Karminfarburg des Glykogens and der Kerne". Z. Wiss. Mikr. , 23:319-22. Bowen , I . D. 1988. "Electron cytochemical localization of acid phosphatase activity in the digestive caeca of the desert locust". J. Roy. Mior. Soc. , 88:279-89. Chiffelle, T.l. and F.A. Putt 1951. "Stain Technique" William and Wilkinco., 26:51. Chou, M.L. 1977. "Anatomy, histology and histochemistry of the alimentary canal of Ghyra mutilata" . J. Herpetol. , 11 (3) : 399-57 . Gomor i , G . 1952. "Microscopic Histochemistry". University of Chicago Press, Chicago. Bulletin Maryland Herpetol og i cal Society Page 131 Volume 20 Number 4 December 1984 Gurr, E. 1962. Halisler, G. 1958. Hayash i , M. , Y 1964. Holt, S . J . 1963. Hugon, J. and 1968. Jennings, J.B. 1962. T962. Kent, P.W. 1971. Loveridge, A. 1955. Mahmound, I.Y. 1979. Mai lory, F. B. 1961 . ' 'Staining Animal Tissue: Practical and Theoretical1 ' . Leonard Hill, London. "Zur Technik und spezifitat des h i stochemi schem Carbanhydrasenachwei ses im Model 1 versuch und in Gewebsschn i tten von Rat tenn i eren . Histochemie. 1 : 29“ 47 • . Nakujima and W.H, Fishman "The cytologic demonstration of $-gl ucuron idase employing naphthol AS - B I glucuronide and hexazonium pa ra rosan i 1 i n ; a preliminary report". J. Histochem. Cytoohem. , 1 2,:293~97. "Some observation of the occurrence and nature of esterases in lysosomes". Ciba Found. Symposium , J.A. Churchill, London . M. Borgers "Fine structural localization of acid and alkaline phosphatase activities in the absorbing cells of the duodenum of rodents". Histochemie. 12:42-66. "Further studies on feeding and digestion in triclad Turbellaria" . Biol. Bull., 123-8. "Hi stochemical study of digestion and digestive enzymes in the rhynchocoel an Lineus ruber". Biol. Bull. 122(1): 63-72. "Biosynthesis of intestinal glycoproteins in animals and man". Gut 1 2 : 4 1 7 • "Reptiles and amphibians from Bahrain and Saudi Arabia". American Documentation Institute No. 4612. Library of Congress, Washington. and T. K1 i cka "Turtles, Perspectives and Research". Ch. 12, A Wiley- Interscience publication, New York. "Pathological Technique". Hafner Publishing Co., New York. Page 132 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December 1984 McCobe, M. and J. Chayen 1965. “The demonstrat ion of la ten particular ami nopept idase activity". J. Roy. Micr . Soc. , 84:361-71. MacManus, J.F.A. and R.W.L. Mowry 1964. “Staining methods". Harper and Row, New York. Mowry, R.W. 1956. “Alcian blue techniques for the h i stochemi cal study and acidic carbohydrates". J. Histoohem. Cytochem. 4:407. _ and C.H. Winkler 1956. “The corolation of acidic carbohydrates of bacteria and fungi in tissue sections with special references to capsules of Cryptocoocus neoformans and Stccphloeoccus . Am. J. Rath., 32:628-29. Novikoff, A.B. 1961. “The Cell". Academic Press, New York. Pearse, A.G.E. 1972. “Histochemistry: Theoretical and Applied". Third edition J. & A. Churchill, London. Re id, R.G.B. 1966. “Digestive tract enzymes in the bivalves Lima hians and My a arenaria". Comp. Bioohem. Physiol. 17: 417“ 33- Rosenbaum, R.M. and B. Ditzion 1963. “Enzymic histochemistry of granular components in digestive gland cells of the Roman snail, Helix pomatia". Biol. Bull. Mar. Biol. Lab. 124:211-24. Smith, E.L. I960. “The enzymes". Academic Press, New York. Suganuma, T., T. Katsuyama, M. Tsukahara, M. Tatematsu, Y. Sakakura and 1981 . F. Murata “Comparative h i stochemi cal study of alimentary tracts with special references to the mucous neck cells of the stomach". Am. J. Anat. 161:214-33. Taib, N.T. 1976. “Anatomy, histology and histochemistry of the alimentary canal of Bentalium entalis L. with some observations on living animals". A thesis submitted to University of Glasgow, U.K., in the candidature for the degree of Philosophiae Doctor. Bulletin Maryland Herpetol og i cal Society Page 133 Volume 20 Number 4 December 1984 Taib, N.T. 1981 . "A h i stochemi cal study of digestion and some digestive enzymes in the lizard Uromastyx aegyptia" . J. Coll. Soi. Univ . Riyadh 12(1) : 1 63” 73 - and B. Jarrar 1983. "Morphology and histology of the alimentary canal of Mauremys caspica Reptilia, Emydidae". Ind. J. Zool. 1 1 (1) : 1-12. 1983. • "Observations on the histochemistry of the alimentary canal of Chaloides levitoni (Reptilia, scincidae)". Arab Gulf J. Scient. Res. 1 ( 1 ) : 1 8 7~ 202 . Tappel , A.L. 1969. "Lysosomes in biology and pathology". North Holland Publ . Co. , London. Wright, R.D., H.W. Florey and A.G. Sanders 1957. "Observations on the gastric mucosa of reptilia". Q. J. Exp. Physiol. Cong. Med. Sei. 42:1-14. Yamada, M. and S. Ofugi 1968. "Silver proteinate method for endopept i dases" . J. Invest. Derrnat. 50:231. Yasuma, A. and T. Ichikawa 1953. "Ninhydrin-schiff and a 1 1 oxan- sch i f f staining, a new h i stochemi ca 1 staining method for protein". J. Lab. Clin. Med. 41:196-99. Department of Zoology 3 College of Science 3 King Saud University } Riyadh , Saudi Arabia Received: 4 April 1984 Accepted: 17 May 1984 Page 134 Bulletin Maryland Herpetol og i ca 1 Society Volume 20 Number 4 December 1984 EFFECTS OF RATTLESNAKE ENVENOMAT I ON UPON AND LABORATORY ( Cvotalus vividis oreganus) MOBILITY OF MALE WILD MICE ( Mus musculus) Abst ract After envenomation by adult rattlesnakes ( Cvotalus vividis oreganus), male wild mice (Mus musculus ) were found to travel significantly farther than male laboratory mice ( M . musculus ) in an open field prior to immo¬ bilization (x = 462 and 206 cm, respectively; range = 0-940 cm). The inequality was attributed to differences in mobility since wild control mice traveled farther during three minutes in an open field than labora¬ tory control mice (x = 1702 and 975 cm, respectively), and latency to immobilization was the same (x = 56 sec). Because wild Mus are probably representative of natural prey items, further assessment of rattlesnake prey trailing ability should include trails of at least 480-500 cm to have ecological validity. Rattlesnakes often rely on an ambush strategy for feeding on rodent prey (Chiszar et al., 1981; Klauber, 1972). Prior to a strike potential prey are detected mostly by visual and thermal cues (Chiszar & Radcliffe, 1976; Klauber, 1972). After striking, adult rodent prey are usually released to prevent damage from rodent teeth and claws, but smaller, less dangerous prey are often not released (Kardong, 1982; O'Connell et al . , 1982; Radcliffe et al., 1 980) . Furthermore, rattlesnake typically do not begin to search for rodent prey until several minutes after a successful strike (Chiszar et al., 1977), apparently to avoid encountering wounded prey still capable of defensive actions (Estep et al . 1981). In an escape response the rodent often wanders several meters before immobilization and death. The snake then locates its dead prey by st r i ke- i nduced chemosensory searching (SICS), characterized by a sustained high tongue-flick rate (see Chiszar & Scudder, 1 980 , and Chiszar et al . , 1982, for reviews). Associated with lateral head movements, SICS allows the snake to contact and follow the chemical trail of the envenom- ated rodent ( Du 1 1 emei jer , 1961; Golan et al., 1982) which, when located, may be distinguished from a nonenvenoma ted carcass (Chiszar et al . , 1980; Duvall et al . , 1978, 1980). Previous studies have demonstrated rattlesnake ability to locate envenomated prey (Du 1 1 erne i jer , 1 96 1 ; Golan et al . , 1 982) , but all experi¬ ments involving this trailing phenomenon have used the laboratory mouse ( Mus musculus). Adequate models of the natural situation would be desirable so that experimental results would better reflect natural predatory behavior. Information concerning prey travel distance after envenomation, necessary for assessing rattlesnake trailing ability, has been provided for the laboratory mouse (Estep et al . , 1 98 1 ) , but not for Bulletin Maryland Herpetol og i cal Society Page 135 Volume 20 Number 4 December 1984 natural prey items. Assuming wild mice ( M . musoulus) are representative of natural prey, the objectives of this study were 1) to determine travel distance of wild mice after envenomat ion, and 2) to compare travel distance following envenomation of wild mice in this study with laboratory mice and data from previous studies. Since wild mice appear to run faster than laboratory mice, it was hypothesized that wild mice travel substan¬ tially farther than laboratory mice after a successful strike. If so, this investigation would indicate that the trailing task confronted by rattlesnakes under natural conditions is greater than previously believed (Estep et al . , 1 98 1 ) . Materials and Methods Eight adult northern Pacific rattlesnakes ( Crotalus vividis oreganus) captured during April 1983 in Walla Walla County, Washington, were maintained collectively in a large (165 x 87 x 86 cm) indoor pen with paper floor coverings, several rocks, and a glass vessel filled with water. All had refused food during semi h i bernat i on (l6-20°C) for three months at an LD 12:12 photoregime. One week prior to start of experimentation the temperature was increased by an electric heater to 25”30°C to induce feeding (Klauber, 1972). Only adult male mice were used in this study to avoid potential sexual differences in mobility (Estep et al., 1981) and venom resistance (Russell, 1980). Male laboratory mice exhibit a greater capacity for each of these measures. Wild mice ( Mus musoulus) were captured from a local barn. Experimental procedures were patterned after a similar study by Estep et al., ( 1 98 1 ) and conducted during January and February, 1984. Snakes were individually placed within a 38-liter aquarium and allowed at least 30 min. adjustment prior to experimentation. Control mice (11 laboratory and 7 wild) were lowered by tongs into the aquarium for 3 ~ 1 0 sec., but held just out of striking range. Each mouse was then immediately transferred to a 100 x 100 x 25 cm open field with 100 10-cm squares. Experimental mice (12 laboratory and 13 wild) were treated similarly except the snake was allowed to strike the mouse after the presentation. The following variables of open field performance were recorded with hand counters and stopwatch: 1) number of squares traversed during each of 12 successive 15-sec. periods (3 min.), 2) latency to immobilization of experimental mice (in seconds), and 3) latency to death (cessation of visible respiration). The open field was wiped clean with a wet towel between trials. Each rattlesnake was allowed to strike and consume at will only one mouse per each of five weeks (6-8 days between tests). Wild and laboratory mice were presented to each snake in a random order. Although predatory and defensive strikes elicit similar effects of envenomation in mice (O'Connell et al . , 1 982) , caution was taken to avoid arousal of defensive behavior in the snakes. Page 136 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December 1984 Resul ts Wild control mice traveled significantly farther than laboratory control mice during the 3 min. in the open field [x = 1702 and 975 cm, respectively; t (11) = 2.54, P < .05; see Figure 1]. Control groups did not differ significantly in weight (x « 20.8 and 22.0 g, respectively). As expected, nonenvenomated mice traveled significantly farther after 3 min. than envenomated mice [F (1,39) - 66.47, P < .001], and the effects of envenomation were realized during the fifth 15~sec. period in the open field (Table 1). More importantly, wild experimental mice traveled significantly farther than laboratory experimental mice [x = 462 and 206 cm, respectively; t (12) = 2.18, P < .05]. Although differences in weight between these two g roups were s i gn i f i cant [x = 19*6 and 22.5 g, respectively; t (19) = 2.1, P < .05], there was no correlation between mouse weight and distance traveled within either group [wild: R (12) = .26, P = .398; lab: R (11) = -.072, P = .824] or both groups combined [R (24) = -.13, P = .538]. Furth ermore, the wild experimental mice in this study appeared to travel farther than laboratory mice of previous studies involving Cvotalus v. vividis (Estep et al., 1981; O'Connell et al., 1982; see Figure 2). That potential interaction involving envenomation vs. nonenvenomat ion and mouse strain was significant in the final 15-sec. period recorded [F (1,39) = 4.25, P - .046; Table 1] does not imply differential susceptab i 1 i ty to envenomation since nearly all envenomated mice were by then immobilized. Variability of the data was substantial . Bulletin Maryland Herpetol og ica 1 Society Page 137 Volume 20 Number 4 December 1984 (WO) 30NV1SIQ 3AllVinwnO X Figure 1 . The cumulative distance traveled by each group in the open field, expressed as number of 10-cm squares entered and centimeters, is shown for each 15-sec. period. Experimental mice were envenomated by a rattlesnake ( Crotalus viridis oreganus) . Page 138 Bulletin Maryland Herpetol og i ca 1 Society SUCCESSIVE 15-SEC PERIODS Volume 20 Number ^ December 1984 w hJ W -d » Ph -O CM O a> *4-1 co o m co . 44 rH 3 (0 0) pci CO CO CM + + ON r- m st • • CM vO ■—* NO •*. m CM vO in CM ON • St -3- r^» oo CM ON VO CO Ov CO CM ON CM • vO O vO ON O O • CM ON + Ov + vO co co ON m o st co ON 00 vO rH CM o • • • • • • • • co CM m rH rH 00 00 m CM 00 CO m 00 + CO CM ON VO vO m o CO CM VO rH vO o o • • o • • • • CM CO 0) a) -a U co v-< 4-1 O 3 I O 4-i bo m 05 + •H »—4 O 00 00 co 4-* m VO O •O Ph •H CM ON CM d co 00 rH m o 0) 'O 4-4 • • • CD • • • • • *h d CD -3- cO CO o O CM CM o rs. o r— 1 *H 4-> Pi rH U rH O vO a. O CD m ex d 4) PH PH H < 5 pH o CD + o -t CM CO r- o vO o ON CO co st CM o st o • • • • • • • • CM o o O o o rH 00 *H i— H vO ON st * r*- 00 *3- CM m o CM o GO rH CM CO rH CM O o • • • • • • • • ON o O 00 o o rH vO 00 st CO CM CM fs. r^ ON o vO o r-- VO co m CM vO o • • • • • • • • o CM vO o CM CO 44 3 > 44 4-i O d d O d d O X w M £ pi M H Bulletin Maryland Herpetol og i ca 1 Society 0) 4-1 c0 0 o d CD % W 4-1 O T3 0) N -O o 4-1 d LAB ESTEP ET AL 1981 9 LAB ESTEP ET AL 1981 ? LAB O'CONNELL ET AL 1982 Figure 2. After envenomation by a rattlesnake ( Crotalus viridis oreganus) , the total distance traveled by mice in this study is compared with previous results involving C. v. viridis. Vertical line = range, horizontal line = mean, rectangle = 1 SD. Page 140 Bulletin Maryland Herpetolog ical Society Volume 20 Number 4 December 1984 DI scusslon The important outcome of this investigation confirmed the hypothe¬ sis that wild mice travel much farther than laboratory mice following rattlesnake envenomat ion. This difference was attributed to the greater mobility exhibited by wild mice in the open field since both suffered immobilization simultaneously. Adjusting for underestimation of the maximal distance traveled by envenomated laboratory mice due to the 2-3 sec. transfer to the open field, Estep et al . (1981) suggested a mean distance of 200 cm would be typical of rattlesnake trailing episodes. Assuming that natural prey items such as Peromyseus and Perognathus (Fitch and Twining, 1946) are capable of similar mobility and are no more susceptible to the effects of envenomat ion, adjustment of the present data suggests a trail length of 480-500 cm might better reflect the ordinary task confronted by rattlesnakes. Neither Peromyseus or Perognathus were available during the time of experimentation (in hiber¬ nation) . However, several mammals including prey items such as Mierotus and Neotoma have recently been found to possess ant '-hemorrag ic resistance to crotalid venom (DeWit, 1982). Accordingly, rodents naturally preyed upon can be expected to travel farther after envenomat ion than mice of similar mobility from populations which suffer no rattlesnake predation. The introduced strain of wild mice used in this study were collected from the latter category. An investigation of the scavenging behavior of western diamondback rattlesnakes found that a rattlesnake might overlook a killed prey item if a second interfering rodent distracted it, later creating a potential situation in which putrefaction would be used as a directional cue for locating lost prey (Gillingham £ Baker, 1981). The apparent selection in rattlesnakes toward holding smaller, nondangerous prey (thereby eliminating the need of trailing) and the data in this study suggest that envenomated prey may also be lost after traveling a distance farther than which rattlesnakes can often negotiate a trail. Perhaps these lost rodents also facilitate scavenging behavior. Leaping behavior of envenomated Peromysous (prev. obs.), which few mice in this study demonstrated, could further complicate the trail. Whether Crotalus viridis exhibits a carrion search strategy remains to be documented. The considerable variability of the data can best be attributed to the method of prey presentation, since the site of fang penetration, which was not controlled, may influence the effects of envenomation (Lester E. Harris, Jr., unpub. data; Kardong, 1982). Because the restrained mice often retaliated with bites directed toward the snake, we often presented the mice in a manner such that the head of the mouse was farthest from the snake. Consequently, many of the mice were struck in posterior regions. Ordinarily, rattlesnakes usually strike prey in the chest or lumbar region (Minton, 1969). Although we did not record the sites of envenomation, it was our impression that rump bites accelerated immobili- Bulletin Maryland Herpetol og i cal Society Page 1 41 Volume 20 Number 4 December 1984 zation of mice, which would seem advantageous in reducing the challenge of trailing prey. However, the risk of an unsuccessful strike launched toward the rear of a forward-moving rodent must be weighed. Hence, it is likely that prey movement, which appears to be significant in releasing the strike of a rattlesnake (Minton, 1969; see also Newman & Hartline, 1982), directs the aim of the strike as well. Since latency for envenomated rodents to die is significantly longer than latency to immobilization, Estep et al . ( 1 981 ) have suggested that strong selective pressures have produced venom components that rapidly inhibit rodent movement whether or not they also contribute to the rodents death. Perhaps selection has also favored venom components contributing to rodent death more strongly in C. v. viridis than in C. v. oreganus. This would be consistent with LD,.,. information for venom toxicity (Glenn & Straight, 1977). In conclusion, it was found that wild mice are more mobile than laboratory mice, and this difference was expressed even after envenomation by a rattlesnake. Moreover, future studies assessing the ability of rattlesnakes ( C . viridis) to follow and locate envenomated rodents should use trails of at least 480-500 cm to have any ecological validity. Literature Cited Chiszar, D., D. Duvall, K. Scudder and C.W. Radcliffe 1983. Simultaneous and successive discriminations between envenomated and nonenvenomated mice by rattlesnakes ( Crotalus durissus and C. viridis) . Behav. Neural Biol. 29:518-521. Chiszar, D. and C.W. Radcliffe 1976. Rate of tongue flicking by rattlesnakes during successive stages of feeding on rodent prey. Bull. Psychon. Soc. 7:485-486. Chiszar, D., C.W. Radcliffe and K.M. Scudder 1977* Analysis of the behavioral sequence emitted by rattle¬ snakes during feeding episodes. I. Striking and chemo- sensory searching. Behav. Biol. 21:418-425. Chiszar, D., C.W. Radcliffe, K.M. Scudder and D. Duvall 1982. St r i ke- i nduced chemosensory searching by rattlesnakes: The role of envenoma t ion- rel a ted chemical cues in the post-strike environment. In D. Mul 1 er-Schwarze and R.M. Silverstein (Eds.) Chemical signals III. Pp 1-24. Chi szar, D. , 1981 . C.W. Radcliffe, H.M. Smith and H. Bashinski Effect of prolonged food deprivation on response to prey odors by rattlesnakes. Herpetol og i ca 37:237-243. Page 1 42 Bulletin Maryland Herpetological Society Volume 20 Number 4 December 19&4 Chiszar, D. 1980 DeWit, C.A. 1982 Dul lemei jer 1961 Duval 1 , D. , 1978 Duval 1 , D. , 1980 Estep, K. , 1981 Fitch, H.S. 1 9^6 G i 1 1 i ngham, 1981 Glenn, J.L. 1977 Golan, L. , 1982 Kardong, K. 1982 and K.M. Scudder Chemosensory searching by rattlesnakes during predatory episodes. In D. Mul 1 er-Schwarze and R.M. Silverstein (Eds.), Chemical signals: Vertebrates and aquatic invertebrates. Pp. 125-139. Plenum Press, New York. Resistance of the prarie vole ( Microtus ochrogaster) and the woodrat ( Neotoma floridana) , in Kansas, to venom of the Osage copperhead ( Agkistrodon contortrix phaeogaster) . Toxicon 20:709-714. P. Some remarks on the feeding behavior of rattlesnakes. Kon. Ned. Akad. Wetensch. Proc. Ser. C. 64:383-396. D. Chiszar, J. Trupiano and C.W. Radcliffe Preference for envenomated rodent prey by rattlesnakes. Bull. Psychon. Soc. 1 1 : 7“8 . K.M. Scudder and D. Chiszar Rattlesnake predatory behavior: Mediation of prey discrimination and release of swallowing by cues arising from envenomated mice. Anim. Behav. 28:674-683. . Poole, C.W. Radcliffe, B. O'Connell and D. Chiszar Distance traveled by mice after envenomation by a rattle¬ snake ( C . viridis) . Bull. Psychon. Soc. 18:108-110. and H. Twining Feeding habits of the Pacific rattlesnake. Copeia 1946: 64-71. J.C. and R.E. Baker Evidence for scavenging behavior in the western diamond- back rattlesnake ( Crotalus atrox) . Z. Tierpsychol. 55: 217-227. and R. Straight The midget faded rattlesnake ( Crotalus viridis concolor) venom: Lethal toxicity and individual variability. Toxicon 15:129-133. . Radcliffe, T. Miller, B. O'Connell and D. Chiszar Trailing behavior in prairie rattlesnakes ( Crotalus viridis). J. Herp. 16:287-293. Comparative study of changes in prey capture behavior of the cottonmouth {Agkistrodon piscivorus) and Egyptian cobra {Naje hade). Copeia 1982: 337“343 . Bulletin Maryland Herpetol og i cal Society Page 143 Volume 20 Number 4 December 1984 Klauber, L.M. 1972. Rattlesnakes - Thei r habi ts, 1 i fe histories, and influence on mankind. Univ. Cal. Press, Berkeley, Calif. Hinton, S.A. 1969. The feeding strike of the timber rattlesnake. J. Herp. 3:121-124. Newman, E.A. and P. H. Hartline 1982. The infrared "vision" of snakes. Sci. Amer. 246:116-127. O'Connell, B., T. Poole, P. Nelson, H.M. Smith and D. Chiszar 1982. Strike- induced chemosensory searching by prairie rattle¬ snakes ( Crotalus v. viridis) after predatory and defensive strikes which made contact with mice ( Mus mus cuius) . Bull. Md. Herp. Soc. 1 8 : 1 - 6 . Radcliffe, C.W., D. Chiszar and B. O'Connell 1980. Effects of prey size on poststrike behavior in rattle¬ snakes ( Crotalus durissus 3 C . enyo3 and C. viridis) . Bull. Psychon. Soc. 16:449-450. Russell, F.E. 1980. Snake venom poisoning. J.B. Lippincott Co., Phi la., PA. — Will lam K. Hayes and Joseph G. Galusha, department of Biological Sciences 3 Walla Walla College 3 College Place3 WA 99324 Received: 11 July 1984 Accepted: 21 July 1984 Page 144 Bulletin Maryland Herpetolog ical Society Volume 20 Number 4 December 1984 EFFECT OF MALE COURTSHIP ON STR I KE- I NDUCED CHEMOSENSORY SEARCHING IN A FEMALE URACOAN RATTLESNAKE ( Crotcdus vegrccndis) AT NATIONAL ZOO Abstract A female Uracoan rattlesnake ( Crotalus vegrandis) at National Zoo exhibited sustained, high rates of tongue flicking after striking rodent prey. During one observation her male cagemate initiated an unsuccessful courtship episode upon seeing the female move. Although male courtship appeared to disrupt her pattern of stri ke- induced chemosensory searching (SICS), she nevertheless persisted with SICS for 1 38 mi n . , compa red with 190 min. four weeks later when no courtship occurred. It is concluded that male courtship interfered with SICS, but only in a quantitative rather than in a qualitative way. Rattlesnakes typically strike and release adult rodent prey (Gans, 1966; Radcliffe et al., 1980), permitting the envenomated rodent to wander up to 700 cm before succumbing to the venom (Estep et al., 1981). Striking rodent prey induces a sustained, high rate of tongue flicking in rattle¬ snakes and other viperids (Chiszar et al., 1977, 1982a, 1983; Dullemeijer, 1981 ; Gillingham & Clark, 1981a). Called st r i ke- i nduced chemosensory searching (SICS), this phenomenon aids the snakes in locating and following the trails left by their envenomated prey (Golan et al., 1982; Chiszar et al., 1983). Tongue flicking transports prey-derived compounds to the vomeronasal organs, and this process contributes to successful trail following (Burghardt, 1970; Burghardt & Pruitt, 1975; Carr et al., 1982; Gillingham & Clark, 1981b; Halpern & Frumin, 1979; Halpern & Kubie, 1980; Kubie & Halpern, .1979; Wilde, 1938). Since rattlesnakes generally do not exhibit much tongue flicking until after they strike prey, SICS has been conceptualized as a transitional component of the predatory sequence of rodent-specializing rattlesnakes, mediating the change from waiting in ambush to searching actively for an ingestible carcass. This transfor¬ mation is relatively long lasting in that SICS endured for an average of 2? hours when prey were removed after envenomation and rattlesnakes were observed until they quit searching (Chiszar et al., 1982b). Most of our research has been conducted with wild-caught rattle¬ snakes that were provided with live rodent prey during captive mainte¬ nance. Rattlesnakes in zoos, on the other hand, are typically fed dead rodent prey, and these snakes rarely have an opportunity to exercise their predatory skills (especially trail following). On-going studies at several zoos are assessing whether or not long-term zoo captives exhibit the same degree of persistence in SICS as wild-caught specimens (O'Connell etal., 1982). The observation here reported was made while measuring duration of SICS at the National Zoo. Bulletin Maryland Herpetol og i ca 1 Society Page 145 Volume 20 Number 4 December 1984 Method Two Uracoan rattlesnakes (Crotalus vegrandis , one male and one female) born at Houston Zoo (5/16/82), and now on exhibit at National Zoo (cage 20), were subjects in the experiment. Each snake had consumed one adult mouse ( Mus musoulusy about 20 g) two weeks prior to the first observation of this study. Also, the pair copulated soon after that feed i ng . The first observation began at 09:00 (2/4/84). All tongue flicks emitted by the female were recorded for 10 successive min. Then a dead mouse (about 20 g) was suspended into the cage from long forceps and held just out of the female's striking range for 3 sec. The mouse was next moved into striking range, whereupon the female struck and released the mouse immediately. The male cagemate was coiled and inactive in the opposite side of the cage, about 50 cm from the female. The mouse was withdrawn after being released by the female, and the observer continued to record tongue flicks until the snake quit searching (defined as 10 successive min. with no tongue flicks). Previous observations of this sort at National Zoo revealed that only the snake that struck prey exhibited SICS; the other snake(s) in the cage usually remained coiled and inactive, even when the searching cagemate crawled near or over them (Chiszar et al., 1984). Hence, there appears to be no social induction of chemosensory searching in rattle¬ snakes. In the present instance the male cagemate did not search for prey, but he initiated an unsuccessful 228 min. courtship episode when the female began to move (about 6 min. after she struck and released the mouse) . One month later, on 3/4/84, the female was again allowed to strike a mouse exactly as described above. The male remained coiled and motion¬ less during the entire time the female exhibited SICS. Comparison of the female's behavior on this second trial with her behavior on 2/4/84 provides an assessment of the effect of male courtship on the female's post-strike activity. Resul ts The female's rates of tongue flicking during the two trials are shown in Table 1. The two. 10-mi n. baseline periods are shown along with the two post-strike periods, each divided into four successive 30-min. sections. It is clear that SICS occurred on both trials and that the female was influenced in only a quantitative manner by the male's sexual overtures during trial 1. Although the male was initially coiled and motionless on 2/4/84, the female's st r i ke- i nduced movements aroused him. He oriented toward her (post-strike min. 6), followed her, and began to emit tongue flicks along her dorsal surface (post-strike min. 11) while she continued with SICS. No qualitative changes in the female's pattern of SICS were obvious to the observers. Page 146 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December lyb^ Table 1 Mean rates of tongue flicking by a female Uracoan rattlesnake observed twice (one month separated the two trials). On each trial a 10-min. baseline (pre-strike) period was recorded along with four successive 30-min. periods after striking a mouse. During the first observation the mal e* cagemate initiated a 228-min. bout of courtship upon detecting the female's first post-strike movements (post-strike min. 6). During the second observation the male remained coiled and inactive for the entire post-strike period. Mean Rates of Tongue Flicking During Successive Periods Date of Observation Basel i ne 10 min. Post- st r i ke min. 1-30 31-60 61-90 91-120 2/4/84 (courtshi p) 0.0 34.7 21.0 6.8 9.9 3/4/84 (no courtship) 0.0 31.9 37.0 22.2 24.8 results of sign tests P>. 05 P>. 05 P<.05 P< . 01 0 • V Q_ On post-strike min. 29 the male wrapped his tail tightly around the base of the female's tail, but the female simply continued with SICS and the male was dislodged after 4 min. by the female's movements. He stayed near her, directing many tongue flicks to her dorsal surface and rubbing his chin over her neck and head. He again wrapped his tail around her's on post-strike min. 40, this time achieving a strong grip which was not broken for 26 min. The female, however, did not respond to the male's courtship by remaining still. I ns tea d,.^, he persisted with SICS, dragging the male about as she moved. The male achieved several more tail wraps later, and several times he used his tail loop to rub the base of the female's tail (Chiszar et al., 1976). After post-strike min. 30 the female's rate of tongue flicking during trial 1 (with courtship) was significantly reduced relative to her rate during trial 2 (without courtship; see Table 1). Although her level of SICS during trial 1 did not return to baseline until post-strike min. 138, her behavior between min. 31 and min. 138 was clearly subdued during the courtship trial. Furthermore, SICS during the second trial did not return to baseline until post-strike min. 190. Therefore, it can be suggested that the male's courtship behavior during trial 1 eventually disrupted the female's searching and hastened its termination. Bulletin Maryland Herpetol og i ca 1 Society Page 147 Volume 20 Number 4 December 1984 Di scussion Courtship by male rattlesnakes has been summarized by Klauber (1956); and Chiszar et al. (1976) reported a peculiar form of tail rubbing that a male massasauga ( Sistvuvus oatenatus tergeminus) used when a female conspecific did not respond to his initial advances. The present male C. vegrandis also exhibited this tail rubbing (first seen on post-strike min. 40) . That the female's level of SICS then dropped sharply may be taken as evidence for a calming effect of this aspect of male courtship. Since the female had mated two weeks prior to trial 1, she was probably in a sexually refractory state during that trial, suggesting that the effectiveness of male behavior in reducing female activity was not dependent upon sexual motivation in the female. Perhaps the main contribution of tail rubbing by the male was simply to mechanically curtail further movement by the female rather than to induce a calm psychological state; but, it is possible that both effects occurred. In any case, SICS was reduced and more quickly eliminated during the courtship trial than during the second trial. Yet it seems remarkable that the female persisted at all beyond post-strike min. 11. The male never discontinued his efforts from this moment through the end of the observation period (min. 138), and he had his tail wrapped around her's for much of this time. Indeed, male courtship continued 90 min. past the termination of SICS by the female, but copulation did not occur. In view of this strong interference by the male, the female's persistence with SICS seems to testify to the obligate nature of this behavior and to its long duration. The purpose of the present observation was to assess the degree of SICS persistence shown by long-term zoo captives, and to compare this with similar measures already taken on wild-caught specimens (Chiszar et al., 1982b). Since most zoos feed dead rodent prey to rattlesnakes, it is generally the case that zoo snakes have lived for long periods without any need or opportunity to execute some aspects of their innate predatory repetoires, especially persistent searching and trail following. Accord¬ ingly, it is possible that predatory behavior may be degraded as a consequence of captive maintenance, and that long-term zoo captives would be at a disadvantage if they were to be released into suitable natural habitat. So far, however, our observations indicate that zoo-maintained rattlesnakes exhibit SICS (O'Connell et al . , 1982), and that SICS persists as long in zoo snakes (including second and third generation captive-born rattlesnakes at San Diego Zoo) as it does in wild-caught specimens that fed exclusively upon live prey in captivity (Chiszar et al., 1982b). The behavior of the present captive-born specimen is consistent with this accumulating data base. Page 148 Bulletin Maryland Herpetolog leal Society Volume 20 Number 4 December 1984 Acknowl edgement We wish to thank James Bacon and Robin Greenlee (San Diego Zoo), Katy Moran (Audubon Park and Zoological Garden), and Dale Marcell ini (National Zoo) for their support of this work. Data on duration of SICS have been gathered at each of these zoos and will be included in a sub¬ sequent publication. Financial support from the M.M. Schmidt Foundation is gratefully acknowledged. Literature Cited Burghardt, G.M. 1970. Chemical perception in reptiles. In J.W. Johnson, Jr., D.G. Moulton and A. Turk (Eds.). Communication by chemical signals , pp. 241-308. Appl eton-Century-Crofts, New York. Burghardt, G.M. 1975. and C.H, Pruitt Role of the tongue and senses in feeding of naive and experienced garter snakes. Physiol. Behav. 14: 1 8 5~ 194. Carr, J., R. Maxion, M. Sharps, D. Weiss, B. O'Connell and D. Chiszar 1982. Predatory behavior in a congenitally alingual russell's viper (Vipera russelli) . 1. Str i ke- i nduced chemo¬ sensory searching. Bull. Md. Herp. Soc. 1 8 : 1 96-204. Chiszar, D., C. Andren, G. Nilson, B. O'Connell, J. S. Mestas, Jr., H.M. Smith and C.W. Radcliffe 1982a. Str i ke- i nduced chemosensory searching in old world vipers and new world pit vipers. Anim. Learn. Behav. 10:121-125. Chiszar, D., C.W. Radcliffe, B. O'Connell and H.M. Smith 1982b. Analysis of the behavioral sequence emitted by rattlesnakes during feeding episodes II. Duration of st r i ke- i nduced chemosensory searching in rattlesnakes ( Crotalus viridis3 C. enyo) . Behav. Neural Biol. 34:261-270. Chiszar, D., C.W. Radcliffe and K.M. Scudder 1977. Analysis of the behavioral sequence emitted by rattlesnakes during feeding episodes. I. Striking and chemosensory searching. Behav. Biol . 21 : 4 1 8-425 . Chiszar, D., C.W. Radcliffe, K.M. Scudder and D. Duvall 1983. Str i ke- i nduced chemosensory searching by rattlesnakes: The role of envenomat ion- rel a ted chemical cues in the post-strike environment. In D. Mu 1 1 er-Schwa rze and R.M. Silverstein (Eds.). Chemical signals in vertebrates Illy pp. 1-23. Plenum Press, New York. Bulletin Maryland Herpetol og i cal Society Page 149 Volume 20 Number *4 December 198*4 Chiszar, D., K.M. Scudder, H.M. Smith and C.W. Radcliffe 1976. An unusual component of courtship behavior in the western massasauga, Sistrurus oatenatus tergeminus . Herpetologica 32:337-338. Chiszar, D., T. Walsh, B. Demeter, M. Davenport, J. Chiszar and H.M. Smith 198*4. No social induction of chemosensory searching in rattle¬ snakes at National Zoo. Abstract of a paper presented at meeting of the Animal Behavior Society, Eastern Washington University, Cheney, Wa. Dul lemei jer, P. 1961. Some remarks on the feeding behavior of rattlesnakes. Ron. Ned . Akad. Wetensoh . Proa . Ser. C. 6*4:383-398. Estep, K., T. Poole, C.W. Radcliffe, B. O'Connell and D. Chiszar 1981. Distance traveled by mice after envenomation by a rattle¬ snake (C. viridis) . Bull. Psyohon . Soc. 18:108-110. Gans, C. 1966. The biting behavior of solenoglyph snakes--its bearing on the pattern of envenomation. Proo . Internat. Symp. Venom. Anim. instituto Butantan, Sao Paulo, Brazil. G i 1 1 i ngham, J 1981. .C. and R.R. Baker Evidence of scavenging behavior in the western diamondback rattlesnake ( Crotalus atrox) . Zeitsohr. Tierphyohol. 55: 217-227. Gillingham, J.C. and D.L. Clark 198la. An analysis of prey searching behavior in the western diamondback rattlesnake, Crotalus atrox. Behav. Neural Biol. 32:235-2*40. Gillingham, J.C. and D.L. Clark 1981b. Snake tongue-flicking: Transfer mechanics to Jacobson's organs. Can. J. Zool. 59:1651-1657. Golan, L., C.W. Radcliffe, T. Miller, B. O'Connell and D. Chiszar 1982. Trailing behavior in prairie rattlesnakes ( Crotalus viridis). J. Herpetol. 16:287-293. Halpern, M. and N. Frumin 1979. Roles of the vomeronasal and olfactory systems in prey attack and feeding in adult garter snakes. Physiol. Behav. 22:1183-1189. Halpern, M. and 1980. J.L. Kubie Chemical access to the vomeronasal Physiol. Behav. 2*4 : 367- 3 7 1 . organs of garter snakes. Page 150 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December 1984 Klauber, L.M. 1956. Ratt1esnakes--thei r habits, life hi stories, and influence on mankind. Univ. Calif. Press, Berkeley. Kubie, J.L. and M. Halpern 1979. Chemical senses involved in garter snake prey trailing. J. Comp. Physiol. Psychol. 93 • 648-667 ♦ O'Connell, B., R. Greenlee, J. Bacon and D. Chiszar 1982. Strike- induced chemosensory searching in old world vipers and new world pit vipers at San Diego Zoo. Zoo Biel. J_: 287-294. Radcliffe, C.W., D. Chiszar and B. O'Connell 1981. Effects of prey size on poststrike behavior in rattlesnake ( Crotalus durissus3 C. enyo3 and 6'. viridis) . Bull. Psychon. Soo. 16:449-450. Wilde, W.S. 1938. The role of Jacobson's organs in the feeding reaction of the common garter snake, Thamnophis sirtalis sirtalis (Linn.). J. Exp. Zool. 77 » 445—465 - — David Chiszar, Psychobiology Program 3 National Science Foundation 3 Washington , D.C. 20550 3 and Dept, of Psychology 3 University of Colorado 3 Boulder 3 CO 80509 ; Joan Chiszar, Armed Forces Institute of Pathology 3 Washington , D.C. 20506; Trooper Walsh and Bela Demeter, Dept, of Herpetology3 National Zoo3 Washington3 D.C. 20008; Hobart M. Smith, Dept, of E.P.O. Biology 3 University of Colorado 3 Boulder 3 CO 80509. Received: 12 July 1984 Accepted: 21 July 1984 Bulletin Maryland Herpetol og i cal Society Page 151 Vol ume 20 Numbe r 4 December 1984 NOTES ON THE ENIGMATIC Barisia imbrioata OF THE BRITISH MUSEUM, AND ON ITS COLLECTION OF REPTILES FROM AMULA, GUERRERO, MEXICO The small collections of reptiles from "Amu la11 (= Almolonga, fide Davis and Dixon, 1959:80), Guerrero, Mexico, obtained by H. H. Smith in 1889, for the British Museum, have long been a source of concern because of the report of Sceloporus variabilis , an Atlantic slope (and trans¬ isthmian Pacific slope) species, and S. aeanthinus , . a t rans- i sthmian Pacific slope species, from there. In order to provide a definitive identification of the enigmatic specimens, and to determine whether locality or previous identification were in error, the entire series was borrowed for study through the courtesy of A. F. Stimson and the authori¬ ties of the British Museum (Natural History). At the same time three enigmatic specimens of Barisia were borrowed to determine whether any might represent species other than B. imbrioata , to which taxon they had long been assigned. The collection purportedly from Amula gives no basis for question¬ ing the veracity of locality data; part of the material had been mi s ident if i ed, and part has never been recorded previously. The examples of Barisia likewise proved to be correctly assigned to B. imbricata3 although two of them exhibit unusual variations. Hence the entire lot of borrowed specimens is reported here to correct and augment the record. Abronia deppei (Wiegmann). The single specimen (1913.7.19.102) is an adult 103 mm s-v, tail 119 mm. It is more or less typical of the species, except that the two lower anterior temporals contact the postoculars, rather than the lower only, and the anterior superciliary contacts the canthol oreal . Otherwise the specimen agrees with descrip¬ tions, having among other characters 26 scales nuchal s to base of tail, six minimum nuchal scale rows, three postoccipi tal scale rows on head, penultimate labial contacting orbital scales, three temporals contacting labials, 3”4 anterior nuchal rows with osteoderms, a small supranasal , a loreal, divided postmental, 119 caudal whorls. The color is excep¬ tionally light although the specimen does not seem to be faded, as the dark crossbars are very distinct. There are six dark crossbars on neck and trunk, nine on tail excluding dark tip. The crossbars are narrow but irregular and broken, separated from each other by spaces almost twice as long as the crossbars. This is the only specimen recorded from regions east of Chilpan- cingo, although Amula is only a short distance (35 km) from the city (see Davis and Dixon, 1959:80, Fig. 1, for localities and vegetational map). Davis and Dixon (1961:82) reported specimens only from the Omiltemi area. Although the specimen has apparently not been recorded previously, it actually must have been available to Boulenger, who in 1913 described Gerrhonotus gadovii from Omiltemi, on the basis of specimens catalogued at the same time as the present example of A. deppei. Page 152 Bulletin Maryland Herpetolog ical Society Volume 20 Number 4 December 1 984 Barisia imbricata imbricata (Wiegmann). Three specimens are of special interest: 71 . 2 . 7 • -4 from "Tehuantepec," Oaxaca, and 1903.9.30.1 18- 119 from "above Xometla," on Mt. Orizaba, 10,000-11,000 ft., Veracruz. The specimen supposedly from Tehuantepec, a female about 94 mm s-v, tail regenerated, has 12 longitudinal dorsal and ventral scale rows, 39 transverse dorsal scale rows, 10-10 supralabial s, 7-8 infralabials, an unpaired postmental, and the mode for all other characters reviewed for the subspecies by Guillette and Smith (1982:19). The pattern is unusual, with dark sides transversed by about 9~ 1 0 narrow vertical light bars; dorsum light tan, sharply delimited from the dark sides along the keels of the 3rd scale row (from midline); a dark brown line down the adjacent halves of the paravertebral scale rows, from occiput onto tail, continuous only on neck, elsewhere broken into sections 2-3 scales long by gaps 0.5“1 scales long; ventral surfaces unmarked. Apparently this specimen is unique among all recorded of the imbricata group, in both pattern (normally unicolor or crossbarred on dorsum as well as sides, never with a median stripe) and the unpaired postmental (always paired). Indeed, these features are characteristic of members of the moreleti group. Nevertheless the aberrant individual is here regarded as simply an anomalous example of B. i. imbricata , with which it agrees in all details of scutellation and body size, whereas it differs in many respects from B. viridiflava , the member of the moreleti group that agrees most closely with the imbricata group. It is not likely a hybrid between 5. imbricata and B. viridiflava , since it has fewer dorsal scale rows (12) than either the former species in Oaxaca (16) or the latter (14). Obviously the locality data are incorrect, but no clue to actual provenience is apparent; presumably the specimen came from somewhere in the southern part of the main plateau of Mexico, definitely not from the Tehuantepec area. One of the two specimens from "above Xometla" (1903. 9. 30. 1 18) is a typical adult, unicolor male 107 mm s-v, with 14 dorsal scale rows. The other specimen, however, a juvenile 62 mm s-v, has 14, 15 or 16 dorsal scale rows at different points on the trunk - the only one with more than 14 recorded since the 1982 review of the species (Guillette and Smith, 1982:19). It also has 42 transverse rows of dorsals, interparietal to base of tail - the maximum recorded for the subspecies (loc. cit.). Presumably these aberrant or unusual character-states have no significance, except as anomalies, since otherwise the specimen is typical of its subspecies . The two specimens demonstrate that B. i. imbricata exists on Mt. Orizaba at altitudes (10,000-11,000 ft.) at least close to if not over¬ lapping with that of the very high altitude B. antauges (12,500 ft. the only reliable record, in Gadow, 1 9 08 : 6 1 ) . The altitude itself is not exceptional, since numerous examples have been recorded elsewhere at altitudes above 10,000 ft. (Guillette and Smith, 1982:30-3), even up to 13,000 ft. on Volcan Popocatepetl (loc. cit.). A more important question, as yet unanswered, is the vertical range of B. antauges ( B . modestus a jr. synonym) on Mt. Orizaba. Bulletin Maryland Herpetol og i ca 1 Society Page 153 Volume 20 Number A December 1 984 Gerrhonotus liooephalus liooephalus Wiegmann. A single example (1913.7.19.103), also catalogued at the same time as the preceding specimen of Abronia deppei and the types of Barisia gadovii (Boulenger), has never been reported in the literature but undoubtedly was a major factor in discovery by Boulenger that his earlier identification of two specimens from Omiltemi of the latter species, reported by Gadow (1905: 195, 233; 1908:380) as Gerrhonotus liooephalus , was in error, particularly since additional specimens of B. gadovii were included in the 1913 collection from Omiltemi. This appears to be a far less common species than B. gadovii in Guerrero; only two reported by Davis and Dixon (1961:53) from Acahuizotla, in addition to the present, are known from the state, whereas dozens of B. gadovii and even numerous Abronia deppei have been reported from the same general area. None has as yet been taken in the Omiltemi area. The Amula specimen is an adult, 1 42 mm s-v, with a regenerated tail; 1A midbody scale rows, 10 on nape; 60 dorsals, interparietal to level of rear margins of thighs; 12 longitudinal rows of ventrals; anterior internasals contacting rostral, separated medially by an irregular-shaped postrostral contacting right posterior internasal but separated from left by an azygous scale; supranasals large but widely separated medially by postrostral; two superimposed postnasals, followed by a loreal superimposed by a canthal; a large cantholoreal narrowly contact i ng frontonasal ; one preocular, entire, contacting anterior superciliary; 2-3 suboculars; 8-8 supralabials to rear edge of subocular, 9th much enlarged, reaching orbit on one side (no postocular separating it); postoculars 1-3; 11-11 supralabials; prefrontals in medial contact; 5-5 inner, 3-3 outer supraocul a rs , 2nd inner broadly contacting prefrontal; frontoparietal s 1-1, separated by frontal-interparietal contact; parietals 1-1, separated posterior to interparietal by a median occipital; anterior and posterior temporals A-A; 9-9 infralabials; postmental asymmetrically divided. Dorsum nearly uniform tan, crossbands very faint; venter unmarked. Although the geographic races of the species are in need of review, it appears that the number of longitudinal rows of dorsals is diagnostic of the nominate subspecies; that character has not been noted in previous comparative accounts (e.g., Tihen, 1 9A8 , 195A; Smith and Taylor, 1950). A revised key to the subspecies, wi th thei r approximate ranges, follows. It seems likely that at least one subspecies has not been named; none has been adequately studied, hence no key can now be regarded as definitive. Key to Subspecies of Gerrhonotus liooephalus 1. Fourteen longitudinal rows of dorsals; a total of three loreal s, canthal s or 1 oreocanthal s on each side; the southern central plateau and Sierra Madre del Sur, Guanajuato, Hidalgo and Queretaro to Guerrero, Oaxaca and western Chiapas - liooephalus Page 15A Bulletin Maryland Herpetological Society Volume 20 Number 4 December 1984 Sixteen or more longitudinal rows of dorsals; loreal -canthal scales 3 or 4 ----- _ - _ _ _ _ _ _ _ _ _ _ _ 2 2. No supranasals; 1 orea 1 -canthal scales three on each side; extreme southwestern Chiapas and presumably adjacent Guatemale - austrinus Supranasals present; loreal -canthal scales four on each side - 3 3. Longitudinal scale rows 18 dorsal ly, 14 ventral ly, and transverse rows of dorsals 55 or more; western Chihuahua - taylori Longitudinal scale rows 16 dorsally o£ 12 ventrally, or transverse rows of dorsals fewer than 55 - 4 4. Second primary temporal usually contacting 5th medial supraocular; southeastern San Luis PotosT - - - * - loweryi Second primary temporal usually separated from 5th medial supra¬ ocular - -5 5. Usually 18 longitudinal dorsal scale rows; southern Sinaloa and adjacent Durango - nomen vacuum Usually 1 6 longitudinal dorsal scale rows - 6 6. Caudal whorls 116-137; tail shorter, 1.75-2.1 times body length; Texas southward to central San Luis PotosT - infemalis Caudal whorls more numerous, 1 40 or more; tail longer, 2.3 times body length; eastern foothills of the Sierra Madre in Veracruz and Puebla - - - - — ophiurus Phrynosoma taurus Duges. Three examples (89.11.13.94-96) are typical of the species; two are adult males, 55“ 59 mm s-v, one a half- grown male 35 mm s-v. All have the ventral scales keeled; femoral pores 12-13, 10-10, 9“ 1 0 respectively. The temporal horns diverge outward in the two smallest specimens, but in the largest curve medially so that they parallel each other toward their tips. Gunther (1890:79) correctly reported these specimens. Only one was taken in the collections reported by Davis and Dixon (1961:43); the species may have become less abundant in recent years. Sceloporus formosus scitulus Smith. A single adult female (89.11.13.93), 63 mm s-v, contains large eggs (10 mm) in the oviducts; 15”15 femoral pores; 31 dorsals; pattern of irregular, small dark spots covering 1-3 scales on back and sides; a pair of dorsolateral light lines on neck, extending posteriorly from orbit, bordered on both sides by a dark line, the lateral one terminating in continuity with a vertical dark line in front of arm insertion; ventral surfaces immaculate. Bulletin Maryland Herpetol og i cal Society Page 155 Volume 20 Number 4 December 1 984 This specimen was reported as S. acanthinus by Boulenger (1897** 497) who noted that Gunther (1890:64) had referred the same specimen to S. spinosus. The taxon has also been recorded from nearby Day i s and Dixon (1961:43-44), at 2.5 mi. S. Almolonga (5600-5800 ft.). The area of intergradation between this and the nominate subspecies remains to be determined. Specimens from Cerro Yucuyagua, 8 km SSE Tlaxiaco, Oaxaca (CUM), represent the nominate subspecies., hence the area of contact with S. f. soitulus must lie somewhere between there and Amula. Sceloporus ochoterenai Smith. Five juveniles (89 . 1 1 . 1 3 • 8 3 ~ 8 7) > 19-24 mm s-v, are readily identifiable with this species by the absence of postrost ral scales (resulting in contact of nasals and internasals with the rostral) , the absence of a postfemoral dermal pocket, and the presence of preanal keels in females. Only two other species of the genus possess the first character, and neither occurs on Pacific slopes of Mex i co . Gunther (1890:75) referred these specimens to S. variabilis , but Boulenger (1897) did not attempt to allocate them, presumably because of their extremely small size. Davis and Dixon (1961:47) reported the species from nearby, at 2.5 mi. S. Almolonga, 5600 ft. Sceloporus spinosus horridus Wiegmann. Five juveniles (89-11 -13- 88-92), the largest 38 mm s-v, the smallest 28 mm s-v, have 3“3(2), 3-4(1), 4-4(2) femoral pores; prefrontal s in contact in three, separated by an azygous scale in one, by contact of frontal with frontonasal in one; and preocular divided in three. These specimens have not previously been reported, although they certainly were available both to Gunther and Boulenger, having been catalogued at the same time as the specimen of S. formosus scitulus. Presumably their small size discouraged any attempt to identify them. They actually are somewhat intermediate between S. s. horridus and S. s. oligoporus, with a higher percentage having fewer than seven femoral pores (40%) than normally expected in 5. s . horridus (19% over-all, fide Smith, 1939:105). The prefrontal s likewise are more frequently in contact (60%) than expected (31%). The preocular character (80% entire on one or both sides) is about as expected, however, for S. s. horridus (82-4) . The significance of the skewness of variation toward norms for S. s. oligoporus apparently lies in the expected geographic range of the latter subspecies a considerable distance up the valley of the Balsas river, to which Amula is fairly close (see map in Davis and Dixon, 1959-80, Fig. 1). A fine series of 15 specimens was reported by Davis and Dixon (1961:45) from Amula, as well as from other localities nearby but on coastal drainages rather than the Balsas drainage. A detailed analysis of variation in these specimens and in other material would clarify the interdigitation of the ranges of S. s. horridus and S. s. oligoporus . It is to be expected that the coastal populations would exhibit little or no influ¬ ence of S . s . oligoporus. Page 156 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December 1984 The concept of conspec i f i c i ty of S. horridus and S. spinosus proposed by Boyer et al . (1982) is here accepted, although evidence to the contrary is in press (Darrell Frost, pers. comm.)* Literature Cited Boyer, Thomas J., Hobart M. Smith and Gustavo Casas-Andreu 1982. The taxonomic relationships of the Mexican lizard species Soeiopovus horridus . Bull. Maryland Herp. Soc., 18(4): 189-191 * Boulenger, George A. 1897. A revision of the lizards of the genus Soeloporus . Proc. Zool . Soc. London, 1897:^74-522, pi. 33* 1913. Descriptions of new lizards in the collection of the British Museum. Ann. Mag. Nat. Hist., (8) 12:583-586. Davis, William B. and James R. Dixon 1959* Snakes of the Chilpancingo region, Mexico. Proc. Biol. Soc. Washington, 74:79“92, figs. 1-2. and 1 961 . Reptiles (exclusive of snakes) of the Chilpancingo region, Mexico. Proc. Biol. Soc. Washington, 74 : 37" 56 , figs. 1-2. Gadow, Hans 1905. The distribution of Mexican amphibians and reptiles. Proc. Zool. Soc. London, 1 905(5) : 1 91 -245, figs. 29“ 32. 1908. Through southern Mexico, being an account of the travels of a naturalist. London, Witherby. xvi, 527 pp., map, ill. Guillette, Louis J., Jr. and Hobart M. Smith 1982. A review of the Mexican lizard Barisia imbricata, and the description of a new subspecies. Trans. Kansas Acad. Sci., 85(1') : 13-33, figs* 1-4. Gunther, Albert C.L.G. I89O. Parts 8-10, pp. 57“ 8 0 , pis. 26-30, in 1885-1902. Biologia cent ral i -amer i cana . London, Porter, xx, 326 pp., 76 pis. Smith, Hobart M. 1939* The Mexican and Central American lizards of the genus Soeiopovus. Zool. Ser. Field Mus. Nat. Hist., 26:1-397, figs. 1-59, pis. 1-31. Bulletin Maryland Herpetol og i ca 1 Society Page 157 Volume 20 Number 4 December 1984 Smith, Hobart M. and Edward H. Taylor . 1950. An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. Bui 1 • U.S. Nat. Mus. , (199) • i-iv, 1-253. Tihen, Joseph 1948. A new Gerrhonotus from San Acad. Sci., 51 (3) : 302-305- Luis Potosf. Trans. Kansas 1554. Gerrhonotine lizards recently added to the American Museum collection, with further revisions of the genus. Am. Mus. Novit., ( 1 687) : 1 - 26 , figs. 1-7. _ Hobart M. Smith, Department of Environmental 3 Population and Organismie Biology 3 University of Colorado 3343 Boulder 3 Colorado 80309 U.S. A. Received: 20 July 1984 Accepted: 1 August 1984 Page 158 Bulletin Maryland Herpetolog ical Society Volume 20 Number k December 1 984 REPTILIAN PARTHENOGENESIS Abst ract Parthenogenet i c species are found in the fishes, amphibians, and in seven families of reptiles. Six parthenogenet i c lizard families are found worldwide in temperate and tropical regions. Parthenogenesis may have arisen separately several times in these families spontaneously or by hybridization. Complete chromosome number may be maintained by pre- meiotic inhibition of mitotic cytokinesis. Invariability of the genetic pool in these species may regulate the habitats they are capable of occupy i ng . In some populations of lizards--in fact, in some entire species of 1 i zards-- there are no males. Females produce offspring without any male input. All of the young are females, genetically identical (except for random mutation) to the mother. They in turn are able to produce young without any contribution from male lizards. This account reviews the mechanisms behind, the significance of, and the frequency with which this phenomenon, known as parthenogenesis, occurs. Many invertebrates may reproduce parthenogenet ical 1 y, but among vertebrates reproduction without fertilization by a spermatozoon is known to occur naturally only in a very few reptiles, amphibians, and teleost fishes. Turkeys can be artificially induced to reproduce pa rthenogene- tically. Fishes with a parthenogenet ic strategy include some perches, darters, and basses. Parthenogenet ic reproduction in amphibians is of a special type known as gynogenesis in which females mate with males and the spermatozoon penetrates but does not fertilize the egg (Cuellar, 1976a). In many vertebrates an- ovum or zygote may be induced to cleave and divide by some physical manipulation such as a pin prick. More interesting is reptilian parthenogenesis in which there is no real mating. In fact, there are no males of the same species with which to mate. Sometimes two females engage in pseudo-copulation; one female having already ovulated acts exactly like a male (except for intromission) and mounts a female with large yolking eggs still in the ovary (Crews and Fitzgerald, 1 980) . But ovulated eggs develop without any external provocation. Although this type of reproduction does not follow the major trend of reproductive evolution in vertebrates and is different from the classic classroom descriptions of vertebrate reproduction, parthenogenesis is neither all that rare nor isolated. One snake and six families of lizards include species which are parthenogenet ic. They occur on all continents in which reptiles are found. The lone species of snake in which parthenogenesis has been dis¬ covered is the Braminy Blind Snake ( Rcanphotyphlops braminus) . It is a small snake of less than 8 inches which burrows among the roots of plants. Bulletin Maryland Herpetol og i cal Society Page 159 Volume 20 Number 4 December 1984 Its distribution is old world including Madagascar, southeastern Asia, East Indies, and many Pacific islands, but it has also been introduced into suitable new world habitats in southern Mexico and Hawaii. Many more lizards are parthenogenet i c , including representatives of the agamids, chameleons, gekkos, lacertids, teiids, and xantusiids (Cole, 1975). On this continent perhaps the most parthenogenet i cal 1 y prolific family, the Teiidae, occurs. Most of these species are found in the deserts of the southwestern United States and northern Mexico. Included are the Cozumel whiptail ( Cnemidophorus cozumela) , the gray checkered whiptail (C. dixoni) , Chihuahuan spotted whiptail (C. exsanguis) , Gila whiptail {C. fZogeZZicaudus) , Laredo striped whiptail (C, Zaredoen- sis) , New Mexican whiptail ( C . neomexicanus) , Sonoran whiptail ( C . opatae) , Yucatan whiptail ((7. rodecki) , Sonoran spotted whiptail ( C . sonorae) , checkered whiptail (C. tes s status) , desert-grassland whiptail (C. uniparens) and the plateau striped whiptail (C. velox) . All of these North American and Mexican parthenogens have arisen in one genus, Cnemidophorus . In South America, teiids which are parthenogenet i c include the spectacled lizard ( GymnophthaZmus undergo o di) , the dwarf teiid {Leposoma percarinatum) , the keeled whiptail (Kentropyx borckianus) and one species that is facultatively parthenogenet i c (that is, it is only parthenogenet ic under certain environmental conditions), the South American whiptail ( C . Zemniscatus) . The lacertid lizards were the first reptiles discovered to be parthenogenet i c (Darevsky, 1958). They are European and occur in the Caucasus Mountains from the Black Sea to the Caspian Sea. They include four species of Wall lizards ( Lacerta armeniaca > L. dahli3 L. vostombekovi3 and L. unisexualis) . The parthenogenet i c geckos are found on many islands of the Pacific and in southeastern Asia. They are the Indo-Pacific gecko ( Hemidactylus garnotii ) and the mourning gecko ( Lepido dactylics Zugubvis) . All lizards so far investigated with three sets of chromosomes are parthenogenet i c. Therefore it is suspected that the triploid gecko ( Gehyra vardegata ogasawarisimae) is also parthenogenet i c. The parthenogenet i c chameleon ( Brookesia spectrum af 'finis) is found in Africa. In Malaya there is a pa r thenogenet i c agamid ( LeioZepis tripZoida) . A facultative parthenogen in Central America is a xantusiid { Lepidophyma fZavimacuZatum) (Cole, 1978). It's obvious from the world wide distribution of pa rthenogenet i c lizards and their occurrence in several families that parthenogenesis has arisen separately several times and that we likely have not discovered all of the pa rthenogenet i c lizard species. It is also possible that, evolving in different taxonomic groups, parthenogenesis came about via different pathways or mechanisms. The mechanism of parthenogenesis, however, must somehow include a form of meiotic restitution. That is, the chromosome number in a haploid egg must at least be restored to the Page 160 Bulletin Maryland Herpetol og i ca 1 Society Volume 20 Number 4 December 1984 full diploid complement equal to that which would have been supplied by both the male and the female (Cuellar, 1971). There are five places during meiosis when this could happen. The first is just before meiosis when the cells in the germinal bed are still dividing mitotically. If during the last mitotic cell division the splitting of the cell membrane and contents (cytokinesis) is inhibited then the divisions of meiosis, which reduce the chromosome number by half, will leave a full complement of chromosomes in the ovum. The second possibility is that during the first chromosome split of meiosis, when homologous pairs (father's matching mother's) line up and separate, this division is aborted. Again there will be twice the chromosome number when the tetrads line up singly and are split at the centromere, during the second meiotic division which reduces the chromosome number by half. Of course, if the second meiotic division is inhibited, and not the first, the ovum will still end up with the diploid (adult) number of chromosomes. The fourth and fifth mechanisms for restoring full chromosome number to an ovum of a parthenogenet i c lizard involves fusion of two nuclei which house the genetic material. During normal meiosis the reduction divisions produce one ovum and 3 inviable polar bodies from one primary oocyte (first stage after germinal bed). Fusion of the nucleus of a polar body with that of the ovum would restore full chromosome number to the ovum. Also if the ovum undergoes cleavage (mitotic division) without fertilization then fusion of the nuclei of the first two cells would result in one cell with a complete chromosome set, which would continue to divide, and one cell with no genetic material. Only one of these mechanisms has any evidence collected in support of it. In the desert-grassland whiptail observations of the chromosomes at the first division revealed a doubled set of chromosomes (Cuellar, 1971). This indicates that this chromosome doubling occurred prior to meiosis. Different investigators argue whether this chromosome restitu¬ tion can occur spontaneously in offspring of bisexual lizards or only in hybrid offspring from a mating of two different bisexual lizard species. The evidence is inconclusive but rather convincing for both hypotheses. In many areas a parthenogenet i c species (e.g., checkered whiptail) is found geographically distributed between two bisexual lizard species (e.g., Western whiptail ( C . tigvis) and the plateau spotted whiptail ( C . gularis septemoittatus)) . The parthenogenet ic lizards are intermediate in ecology and morphology as well as distribution between their bisexual neighbors. The New Mexican whiptail has been found to be cytol og i cal 1 y and biochemically intermediate between the Western whiptail and the little striped whiptail ( C . inornatus) (Brown and Wright, 1979). All of these factors point toward hybridization as the trigger for the unusual chromo¬ some restitution. But there are cases in which the lizard is facultatively parthenogenet ic and/or there are no similar (congeneric) bisexual lizards living in habitats adjacent to the parthenogenet i c species (e.g., C. lemnisoatus and L. flavimaoulatum) . In these cases there would be no stock from which to derive a parthenogenet ic hybrid. Even more convincing support for the idea of spontaneous chromosome restitution is the evidence Bulletin Maryland Herpetol og i cal Society Page 1 6 1 Volume 20 Number 4 December 1984 from intraclonal histocompatibility studies. If a piece of skin is grafted from one lizard to another It should only be accepted by the second lizard if it is essentially genetically identical to the first. There appears to be remarkable genetic integrity between individuals of a parthenogene- tic species (Cuellar, 1976b). The degree of genetic homogeneity suggests that an entire species may have evolved from one individual lizard. It is likely that hybridization would occur infrequently but often enough to contribute more heterogeneity to the gene pool of the parthenogenetic species. Whatever the mechanisms responsible, the fact that parthenogenetic species do exist represents an unusual opportunity to look into the advantages associated with sexual reproduction and nonsexual reproduction. The parthenogenetic lizards obviously have the advantage of complete reproductive capacity in every animal rather than reproductive capacity being attained only through union of two animals. More simply, all parthenogenetic lizards are females so they can all produce eggs. Normal sexual lizards have populations which are only half female, therefore only half the population can produce eggs. If you assume that the partheno¬ genetic lizards must be ecologically very similar to the sexual lizards from which they sprang then it is very difficult to understand why there are any similar sexual lizards left. Imagine a population of 12 sexual lizards and one newly derived parthenogenetic lizard. For simplicity, let's assume that the clutch size of these animals is two and that the adults die when the young are hatched — even replacement. In the next generation there would remain 12 sexual lizards (2 from each of six females) but there would now be 2 partheno¬ genetic 1 izards, aga in both females. The succeeding generations would see populations with 4, 8, 16, 32, and 64 parthenogenetic lizards in the 7th generation, but there would always be only 12 sexual lizards. Remembering that the parthenogenetic lizards have the same genes as the sexual lizards from which they came, they are subject to the same ecological limitations as the sexual lizards. If a natural event decimated the population or if many of the lizards died because carrying capacity of the habitat had been surpassed, a proportionate number of sexual and parthenogenetic lizards would die. If a natural event in the seventh generation reduced the population size back to twelve, eleven of the lizards would now be parthenogenetic. Since it takes two sexual lizards to produce any offspring it's obvious that the following genera¬ tion would contain only parthenogenetic lizards. But this does not occur. Sexual lizards remain dominant in their habitats and parthenogenetic lizards often are found only on the periphery of the habitat, or in habitats which are not stable, such as river washes (Cuellar, 1977; Wright and Lowe, 1968). This may suggest that the variability between sexual individuals allows each animal to exploit resources different enough from those of his neighbor that these animals don't always have to compete. Since parthenogenetic 1 izards are cl ones Page 162 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December 1984 with no variability (except that due to random mutation), competition between individuals may be very much greater. This competition may reduce the energy stores needed for reproduction, establish stress- i nduced hormonal inhibition of reproduction, or both. Parthenogenet i c lizards represent a natural experiment of the evolutionary trend toward greater genetic recombination. Discovering and exposing new parthenogenetic species and their habitats may help give some insight into the conditions which favor sexual differentiation by looking at those which do not. Paradoxically they may help us understand the basic principles common to most vertebrate species, that is sexual species . Acknowl edgment I would like to thank Hobart M. Smith for suggestions on and critique of this manuscript. Literature Cited Brown, W.M. and J.W. Wright 1979. Mitochondrial DNA analyses and the origin and relative age of parthenogenetic lizards (genus Cnemidophorus) . Science 203:1247-1249. Cole, C.J. 1975. Evolution of parthenogenetic species of reptiles. In "Symposium on Intersexuality in the Animal Kingdom" R. Reinboth, ed., Spr i nger-Verl ag pp. 340-353. 1978. The value of virgin birth. Nat. Hist. 87:58-63. Crews, D. and K.T. Fitzgerald 1980. "Sexual" behavior in parthenogenetic lizards ( Cnemido - phorus) . Proc. Natl. Acad. Sci. USA 77 :499“ 502. Cuellar, 0. 1971. Reproduction and the mechanism of meiotic restitution in the parthenogenetic lizard ( Cnemidophorus uniparens) . J. Morph. 133:139-166. T976a. Cytology of meiosis in the triploid gynogenetic salamander Amby stoma tvembtayd. .Chromosoma 58:355-364. T97£b. Intraclonal histocompatibility in a parthenogenetic lizard. Evidence of genetic homogeneity. Science 193: 150-153. Bulletin Maryland Herpetol og i cal Society Page 1 63 Volume 20 Number 4 December 1984 Cuellar, 0. 1977* Animal parthenogenesis. A new evolutionary-ecological model is needed. Science 197:837-843. Darevsky, I.S. 1958. Natural parthenogenesis in certain subspecies of rock lizards, Laoerta saxioola. Dok. Akad. Nauk. S.S.S.R. 122:730-732. (English trans. publ . by Am. Inst. Biol. Sci., 877-879). Wright, J.W. and C.H. Lowe 1968. Weeds, polyploids, parthenogenesis, and the geographical and ecological distribution of all-female species of Cnemidophorus . Cope i a 1968:128-138. — Cl iff H. Summers, Laboratory of Comparative Reproduction , Department of Environmental, Population, and Organismic Biology , Box 324, University of Colorado, Boulder, CO 80309 Received: 9 June 1984 Accepted: 21 July 1984 Bulletin Maryland Herpetol og i cal Society Page 164 Volume 20 Number 4 December 1984 A NEW SUBSPECIES OF Centrolenella orientalis (ANURA: CENTROLEN I DAE) FROM TOBAGO, WEST INDIES Rivero (1968) described Centrolenella orientalis on the basis of a single specimen collected on Mt. Turumiquire, Venezuela, June 24, 1967 (MCZ 72497). A strikingly similar frog (Fig. 1) was recently discovered on the island of Tobago, West Indies (Hardy, 1977). Duellman (1977), working without benefit of specimens or field experience, assumed that the Tobago Centrolenella was identical to Centrolenella orientalis of Venezuela and published the rather remarkable statement that Centrolenella orientalis occurs in "the mountains of northeastern Venezuela" (thus implying a broader geographic range than was previously known) and on the island of Tobago. Zweifel (1977) noted that the distance between Tobago and Mt. Turumiquire is 370 kilometers, and pointed out that Duellman's comments should have been documented. In 1982 I listed the Tobago frog as Centrolenella cf orientalis , primarily because I had been unable to collect additional specimens of Centrolenella orientalis in Venezuela. At the present time Centrolenella orientalis remains known only from the holotype. Recent work in Tobago, however, coupled with a detailed study of morphometric data from both populations, suggest that the Tobago frog is, indeed, conspecific with Centrolenella orientalis of Venezuela, but is subspeci f i cal 1 y distinct. The Tobago population shall be known as: Centrolenella orientalis tobagoensls subsp. nov. Holotype. USNM 195045, collectedon August 31, 1972, along the Roxborough-Pa rtatuv i er Road in the vicinity of Bloody Bay, St. John Parish, Tobago (Fig. 2) . Paratypes. Twenty-nine specimens from Tobago as follows: USNM 192745, Windward Road, vicinity of mile marker 22? near Lambeau Hill Crown Trace, St. Paul Parish, 11 July 1971; USNM 194999-5000, Windward Road, vicinity of mile marker 22?, near Lambeau Hill Crown Trace, St. Paul Parish, 28 August 1972; USNM 195031-34, Windward Road, vicinity of mile marker 22^, near Lambeau Hill Crown Trace, St. Paul Parish, 30 August 1972; USNM 195039, Charlottevi 1 1 e-Bloody Bay Road, near Hermitage, vicinity of mile marker 30, St. John Parish, 30 August 1972; USNM 195040, Charlottevi 1 1 e-Bloody Bay Road, near Hermitage, vicinity of mile marker 27?, St. John Parish, 30 August 1972; USNM 195044, Roxborough-Partatuv ier Road, vicinity of Bloody Bay, St. John Parish, 31 August 1972; USNM 195152-55, Windward Road, vicinity of mile marker 22?, near Lambeau Hill Crown Trace, St. Paul Parish, 14 September 1972; USNM 195157, Windward Road, vicinity of mile marker 22?, near Lambeau Hill Crown Trace, St. Paul Parish, 16 September 1972; USNM 227732, 3.75 mile WSW of Charlottevi 1 le, Northside (Bloody Bay) Road, mile Bulletin Maryland Herpetol og i cal Society Page 1 65 Volume 20 Number 4 December 1984 Figure 1. The Tobago Centrolenella photographed alive. A. Dorso- 1 atera 1 view. B. Ventral view. Page 1 66 Bulletin Maryland Herpetolog ical Society Volume 20 Number 4 December 1984 marker 27i, St. John Parish, 17 November 1971; USNM 227733-40, 3.75 miles WSW of Charlottevi 1 le, Northside (Bloody Bay) Road, mile marker 27i, St John Parish, 16 November 1971; USNM 227741-42, Windward Road, vicinity of mile marker 25i, near Lambeau Hill Crown Trace, St. Paul Parish, 16 November 1971; USNM 227743, 2 miles WSW of Chari ottev i 1 1 e, Northside (Bloody Bay) Road, Hermitage, at mile marker 29*5, St. John Parish, 16 December 1978; USNM 227744-45, 2 miles WSW of Charlottevi 1 le, Hermitage Bridge, along Northside (Bloody Bay) Road, at mile marker 29i, 23 December 1978. Range. Known only from mountainous areas on the island of Tobago, West Indies (Fig. 3) . Diagnosis. Maximum length 22.9 mm; tympanum hidden; vomerine teeth and humeral spines absent; first finger equal to or longer than second; toes three-fourths webbed; dorsal surfaces green, with or without black or yel low punctations; belly granular; dorsal surfaces smooth. Description of the holotype. A male; snout-vent length, 21.7 mm; head width, 7.9 mm; tibia length, 10.8 mm. In preservation, dorsum light straw-colored and with about twelve conspicuous black dots scattered randomly among a pattern of evenly-spaced, small mel anophores . Similar small mel anophores in a narrow band along the top of the femur, and continuing down the leg to the bases of the toes. Upper arm unpigmented throughout. Lower arm with scattered melanophores. Ventral surfaces flesh-colored, un-pigmented. Color in life (based on recent field observations). Body pale leaf-green above, with or without scattered small black melanophores, larger black spots, and lemon-yellow punctations. Upper jaw flesh-colored, unpigmented. Limbs, except upper arm, green dorsal ly, with or without punctations. Upper arm flesh-colored, entirely unpigmented. Femur with a narrow line of green dorsal ly, otherwise flesh-colored. Toes and toe pads pale to deep lemon yellow. Belly transparent centrally, whitish or pale green and somewhat opaque toward the edges. Throat flesh-colored, or extremely pale green. Eye bright canary yellow with scattered, large melanophores which tend to form a ring around the pupil. Variations in the paratypes. The paratypes vary from 20.2 to 22.9 mm in snout-vent length, and from 10.5 to 12.2 mm in tibia length. In the preserved series, large dorsal pigment spots may be present or absent (Fig. 4); otherwise the paratypes are remarkably similar to one another. Compar i sons. Centvolenelta orientalis tobagoensis may be distin¬ guished from Centro Zene t la oT'Lenta'l'is orienta'l'is on the basis of relative tibia length (shorter in Tobago), and, presumedly, less intense pigmenta¬ tion. Bulletin Maryland Herpetolog i cal Society Page 1 67 Volume 20 Number 4 December 1984 Figure 2. Centrolenella orientalis tobagoensis , the holotype (USNM 195045). A. Dorsal view. B. Ventral view. Page 1 68 Bulletin Maryland Herpetol og i ca 1 Society Volume 20 Number 4 December 1984 TOL AGO F?9ure 3. Distribution of Centrolenella orientalis orientalis and Centrolenella orientalis tobagoensis in northeastern Venezuela and the southeastern Caribbean. Rivero (1968) gives a snout-vent length of 21.2 mm, and a tibia length of 12.2 mm for the holotype of Centrolenella orientalis orientalis. These figures result in a snout-vent/tibia length ratio of 1.74. I have examined the holotype of Centrolenella orientalis orientalis and agree with Rivero's original measurements. In Centrolenella orientalis tobagoensis snout-vent/tibia length ratios vary from 1.84 to 2.04 with a mean of 1.93 (Fig. 5) . Results of statistical analysis done on these ratios indicate that Centrolenella orientalis tobagoensis is morphometrically distinct from Centrolenella orientalis orientalis . The standard deviation of the Tobago population is 0.056. The 1.74 snout-vent/tibia length ratio of the single specimen of Centrolenella orientalis orientalis is 3.4 standard deviations less than the mean value of this ratio for Centrolenella orientalis tobagoensis . Assuming that these ratios are normally distributed in Centrolenella orientalis tobagoensis , the probability of observing a value as low as 1.74 in this population is 0.0003 or 1 in 3,333- Application of a statistical test for comparing a single observation to a sample mean (Sokal and Rohlf, 1969) resulted in a "t" value of -3*338 which is significant at the a = 0.01 level. Rivero (1968) stated that Centrolenella orientalis orientalis is green below, tending toward blue on the throat and chest, and that the discs of the toes are somewhat orange. In recently collected specimens from Tobago, the ventral surfaces of the body are extremely transparent, Bulletin Maryland Herpetolog i cal Society Page 1 69 Volume 20 Number 4 December 1984 Figure 4. Paratypes of Centrolenella orientalis tobagoensis showing variation in dorsal pigment. A. USNM 227737. Large dorsal spots entirely absent. B,C. USNM 227736, USNM 195152. Moderately developed dorsal spots (typical of approximately 90% of the paratypes). D. USNM 195155. Maximum development of dorsal spots. This single specimen approaches the condition seen in the holotype of Centrolenella orientalis orientalis. Page 170 Bulletin Maryland Herpetol og i ca 1 Society Volume 20 Number 4 December 1984 c. oriental's tobagoensis C. orientals orientalis 1.72 1.76 1.80 J I 1 I I I - 1 — - 1 - 1 - 1 - L 1.84 1.88 1-92 1-96 2 00 2-04 Figure 5. Statistical analysis of the snout-vent/tibia length ratios ~ of the two populations of Centrolenella orientalis. The broad band represents one standard deviation on each side of the mean. becoming somewhat opaque toward the sides, and may be pale flesh-colored, whitish, or very pale light green. The throat and chest regions are flesh-colored or pale transparent green, and there is no suggestion of blue pigment. The toes and toe discs are yel low throughout. Tiny yellow punctations occur on the dorsal surfaces of at least some specimens of Centrolenella orientalis tobagoensis. Similar punctations were not mentioned in the type description of Centrolenella orientalis orientalis. The holotype of Centrolenella orientalis orientalis has numerous large, black spots over most of the head and body. Only one specimen of Centrolenella orientalis tobagoensis approaches this pattern (USNM 195155, see Fig. 4). If the holotype of Centrolenella orientalis orientalis is assumed to be an average specimen, then Centrolenella orientalis tobagoensis is probably a less well-spotted frog. As Rivero (1968) pointed out, it is unfortunate that the eye pigment of Centrolenella orientalis orientalis has not been described. D i scussion. The northeastern limits of the range of the family Centrol enidae in Venezuela is not well known. Rivero (1980) indicates the northeastern edge of range as the vicinity of Mt. Turumiquire. Juan Leon (personal communication) states, on the other hand, that the genus Centrolenella occurs on the Paria Peninsula. Regardless of the range of Centrolenella in Venezuela, there is good evidence to show that this genus does not occur on the island of Trinidad (Kenny 1969, 1977). The Bulletin Maryland Herpetol og ical Society Page 171 Volume 20 Number 4 December 1984 Tobago Centrolenella is, in fact, one of a number of vertebrate animals (4 frogs, 1 lizard, 1 snake, 11 birds, and 2 mammals) which occur on Tobago but not in Trinidad and are striking.ly similar or identical to species occurring in Venezuela (Hardy 1977, 1982, 1983). The biogeogra- phical implications of these distributions, and the exact relationship of Centrolenella orientalis tobagoensis to Centrolenella orientalis orientalis will not be understood until more material is available from Venezuela and until biochemical differences (or similarities) between these various disjunct populations have been documented and critically analyzed. Acknowl edgments Work on this project was initially supported by a grant from the American Philosophical Society, and subsequent field work was supported by a fellowship from the Organization of American States, and a grant from the Center for Field Research (Earthwatch) . Dr. George Drewry, of the Office of Endangered Species, U.S. Department of Interior; Mr. Robert Tuck, Jr., of Cero Cosa Community College, California; and Janet Olmon, formerly of the Virginia Institute of Marine Science, provided useful assistance in the field. Dr. Kenneth Turgeon, NESDIS, National Oceano¬ graphic and Atmospheric Admi n i strat ion, ass i sted with the statistical analysis, and Sherry Picciolo provided a precise translation of the type description of Centrolenella orientalis orientalis. L i terature C ? ted Duel 1 man , William E . 1977* Das Tierreich. Lieferung 95- Liste der rezenten Amphibien und Reptilien, Hylidae, Cent rol en i dae, Pseudidae. Wal ter de Gruyter , Berlin, xix + 225p. Hardy, Jerry D. , Jr. 1977. Frogs, islands and evolution (abstract), p. 5, J_N Eastern Seaboard Herpetol og i cal League. Program and Abstracts of Papers. 10th Biannual Meeting, 5 March 1977, 1 2p . • 1 982. Biogeography of Tobago, West Indies, with special reference to amphibians and reptiles: A review. Bull. Md. Herp. Soc. 1 8 (2) : 37" 1 42 . 1W7 A new frog of the genus Colostethus from the island of Tobago, West Indies (Anura: Dendrobat i dae) . Bull. Md . Herp. Soc. 19(2) : 47-57- Page 172 Bulletin Maryland Herpetol og i ca 1 Society Volume 20 Number 4 December 1984 Kenny, Jul ian S. 1969. The amphibians of Trinidad. Stud i es on the Fauna of Curacao and Other Caribbean Islands 29Tl08):1-78 + 15 plates . 1977. The amphibia of Trinidad: An addendum. Stud i es on the Fauna of Curacao and Other Caribbean Islands 51 (1^9) :92- 95. Rivero, Juan 1968. A. Los Cent rol en i dos de Venezuela. Memoria de la Sociedad de Ciencias Naturales La Salle de Venezuela. Carl bbean J. Science 4(1) :397_405. 1980. Anfibios neot rop i cal es : Origen y Distribucion, J_N Salinas Pedro J. [editor]. Zool . Trop. Actas, Congreso Latin. Zool . 2:91-123. Sokal , Robert R. and F. James Rohlf 1969. Biometry. The principles and practice of statistics in biological research. W. J1. Freeman and Co., San Francisco xx i + 77&p. Zweifel , Richard G. 1977. Liste der rezenten amphibien und reptilien - Hylidae, Centrol enidae, Pseudidae, by William E. Duellman, 1977. Das Tierreich, Lief. 95, xix + 225p. 280 DM (book review). Herpetological Reviews 8(3) : 8 1 —83 . — Jerry D. Hardy, Jr., Department of Herpetology, Natural History Society of Maryland, Inc,, 2643 North Charles Street, Baltimore, Maryland 21218. Received: 27 August 1984 Accepted: 12 September 1984 Bulletin Maryland Herpetological Society Page 173 Volume 20 Number 4 December 1984 NEWS AND NOTES: THE HERPETOLOGICAL WORKS OF JACOB KLEIN, WITH EMPHASIS ON THEIR PERTINENCE TO MEXICAN HERPETOLOGY Abstract The works of Jacob Theodore Klein are reviewed with attention to their contribution to the field of herpetology In general and Mexican herpetology in particular. With the exception of coinage of the word herpetology (although defined differently from at present), his labors now seem an exercise in futility, although they undoubtedly served a useful purpose in their own era in channelling the efforts of others in more fruitful directions. Jacob Theodore Klein ( 1 68 5“ 1 7 59) is today little known, at least in the Americas, yet was one of the most prolific naturalists of his era He wrote at least 24 different books on diverse organisms, appearing in a total of at least 31 different editions and printings. Separate books were published on botany, echinoderms, marine worms, molluscs, inverte¬ brate fossils, geology, fishes, birds, limbless tetrapods, quadrupeds and reviews of Linnaean works, most of them synoptic of then current knowledge. It was an amazing output for his time and for leisure-hour pursuit, for as Secretary of the State of Danzig he had little spare time. The seven works pertaining to amphibians and reptiles appeared between 1743 and 1760 (see Literature Cited). They were taxonomic in nature, listing species then known, or their classification, or both, with brief characterization. Two works of 1760 are post 1 innaean , yet not a single name is now attributed to Klein, and his works served in no way as a source for others. The primary reasons for the total lack of nomenclatural impact of his publications are (l) the almost exclusively compilatory nature of his writings, summarizing and to a certain extent analyzing the works of others, lacking any "primary" contribution what¬ ever, and making only "secondary" innovations in arrangement and group names for species recognized by others (although he apparently did have a museum of his own, as indicated by the portrait serving as a frontis** piece for his Systeme Naturel of 1754, showing specimens, including some bottled snakes and other herps, of various sorts); (2) the misfortune of being prel innaean except for two posthumous works of 1760; (3) the failure to adopt consistently the principles of binomial nomenclature (required for any work officially admissible in biological nomenclature) and (4) the inconsistency of cl ass i f icatory schemes adopted in his several works. Page 174 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December 1984 Indeed, perhaps at present Klein's greatest claim to fame in herpetology is his creation of that very word in his 1755 work, even though his "herpeta" (i.e., herpetozoans) embraced all limbless animals then known that have an elongate body and move sinuously (1755:1)- That definition limited the coverage to snakes, "amph i sbaen i ds" (including caecilians) and worms - the extremes of “creeping and crawling things," as implied etymologically. He thus thought of “herpeta" as the antithesis of “quadrupeda" or four-1 imbed animals, which he interpreted quite literally, not in the present “tetrapod" sense including all members of the classes Amphibia, Reptilia, Aves and Mammalia. With the one exception, then, of creation of the word “herpetology," Klein's contri but ions are now lamentably of historical interest only. They are nevertheless a monument to industry and human frailty, and in their time undoubtedly served a useful purpose in exploring a trail whose destiny could not be anticipated but which in its dead end showed that the goal had to be reached some other way. Trial and error has always been the rule when the goal is clear but the route to it is unclear. Those who choose blind alleys often deserve as much homage as those who guess aright, for neither knows how their efforts will succeed; failures are often as important as successes, when learning results. An over-all view of Klein's publications, from the earliest to the latest, gives a clear picture of the heroic struggle early workers endured in striving for the most “natural," meaningful classification of organisms. Klein in the end was convinced that superficial appearance and habitat were proper criteria for assumption of relationship - a perfectly reason¬ able conclusion on the premise of special creation that was then prevalent. Relationship through evolution and phylogeny was then dimly comprehended, or not at all; our present classification could stand on no other basis. Linnaeus suffered from the same misconception, but came closer to a true “natural" classification than Klein because of greater attention to details of structural similarity. Klein's struggle in grappling with classification is evident also in the extensive inconsistency in application of names in various cate¬ gories, from the highest to the lowest. Indeed, often names as such were not used, placing reliance instead upon a description. Generic names were not used throughout given genera, or different ones were used for the name in some cases. Species were not always given specific names, or even a brief polynomial, and sometimes were not even given the generic name. The idea of binomial nomenclature had not yet been adopted, and indeed all names and ranks seemed to be rather nebulous. (Thus the assignment of Klein's names to given ranks in the following accounts takes some liberties with his actual usages). Despite these shortcomings in Klein's work, as seen in retrospect, we here seek belatedly to promote some of the attention that Klein's nobl e efforts deserve, by briefly reviewing all of his herpetol og i ca 1 works of which we are aware, noting in the process their pertinence to Mexican herpetology. Bulletin Maryland Herpetol og i ca 1 Society Page 175 Volume 20 Number 4 December 1984 1743 The earliest work pertaining to herpetology appeared in 1743 and was a tentative review, in Latin, of the higher categories of quadruped (the term being used in its literal sense, in reference strictly to four- footed animals) classification. The "Class Amphibia," embracing all amphibians and reptiles (pp. 25“30) , was diagnosed, and two orders, Reptilia (with three "genera:" Testudo^ Rana and Laoerta) and Serpentia (with one genus, Anguts) , were all briefly discussed with passing mention of a few species and pertinent literature. 1751 The second work (1751:98-123) also is limited to quadrupeds, but with greater detail, listing all known species-group taxa. Again, "quadruped" is used in its literal sense, hence excluding snakes and other limbless tetrapods. The "Order Depilata" contained three divisions (rank not specified): Testudinata, Cataphracta and Nuda. All turtles (Testudinata) were placed in Testudo , a group presumably equivalent to a genus although the rank was not specified. The species of Testudo were placed in two groups-one with fused digits ("pedibus anomalis"), containing the sea turtles (three species), the other with digits distinct ("digitis discretis"), containing eleven species. One of the latter was presumably Mexican: " Testudo ex nova Hdspanda, sourced from "Seba, p. 129, pi. 80, fig. 5*“ The crocodilians were subsumed under the "Cataphracta," and perhaps under one genus, "Croooditus ," but the species were poorly de¬ limited; a Croooditus amerioanus was recognized, another stated to be the alligator, and " caimanos " of diverse origins (Ceylon, Africa, America). The "Nuda" embraced nine genera. Laoerta, by far the largest, contained 88 species, arranged in three groups (crestless, crested, salamander-1 ike) . At least twelve may be Mexican: (1) Laoerta major y supposedly the same as Hernandez1 (1648) "ti lquetzpal 1 in," sourced from "Seba, p. 152, pi. 97, fig. 2" (Smith, 1989:9, identified Hernandez' species as Soeloporus aeneus and S. grammious) ; (2) Laoerta texixinooyotl, with a name derived from Hernandez (1648), although not mentioned, but sourced from "Seba, p. 151, pi . 98, figs. 1-3“ (Smith 1969:9, suggested that Hernandez1 species might be S. jarrovi) ; (3) Laoerta de Taleteoy equated with Hernandez' (1648) "tamacolin" (regarded by Smith, 1969:9, as any of several common species of anurans of the genera Eleutherodao- tylus j Leptodaotylus , Hyla, Soaphiopus and Bufo)y sourced from "Seba, p. 151, pi. 97, fig. 1; (4) Laoerta Brazily included here only because it is given an Aztec name (quetzpaleo, not mentioned by Smith, 1969; perhaps a misspelling of "cuetzpal 1 in," see no. 5), although without reference to Hernandez, sourced from "Seba, p. 152, pi. 97, fig. 4;" (5) Laoerta Mexioana Cutezpallin (=cuetzpal 1 in of Hernandez, 1648, but not mentioned; regarded as some species of Soetoporus by Smith, 1969:9) , Page 176 Bulletin Maryland Herpetological Society Volume 20 Number 4 December 1984 sourced from "Seba, p. 152, pi. 97, fig. 5;" (6) Laoerta Mexioana, sourced from "Seba, p. 31, pi . 30, fig. 2:" (7) Laoertus Amerioanus3 noted as one of the "iguanas," may be a Mexican species, sourced from "Seba, p. 149, pi. 95, fig. 1," (8) Laoerta Mexioana3 sourced from "Seba, p. 140, pi. 89, fig. 1;" (9) Laoerta Mexioana3 called "tecoixin" (= tecuixin of Hernandez, 1648, not mentioned, but regarded by Smith, 1969: 9, as any of several small species of Soeloporus or as Urosaurus bioarinatus) , sourced from "Seba, p. 1 4 1 , pi. 89, fig. 2;" (10) Laoerta he liaca3 Americana 3 peotinata 3 may be Mexican (no indication of locality), sourced from "Seba, vo 1 . 2, p. 169, pi. 106, fig. 2;" (11) Laoerta Salamandrina3 Salamandra Mexioana 3 sourced from "Seba, p. 21, pi. 20, fig. 4;" and (12) Laoerta Tapayakin3 or Laoerta orbicularis (= tapayaxin of Hernandez, 1648, not mentioned; Phrynosoma orbioulare as long accepted, and as reiterated by Smith, 1969: 10), sourced from "Seba, vol . 2, p. 10, pi. 8, fig. 7." Other genera recognized among Klein's "Nuda" were Salamandra 3 Gekko3 Cordylus3 Soinous3 Seps3 Chamaeleo3 Rana and Bufo3 among which only three species are possibly Mexican: (l) Cordylus cauda bifuroata3 Salamandra Amerioana3 sourced from "Seba, vol. 1, p. 173, pi . 109, fig. 5;" (2) Chamaeleo Mexioanus3 or "Cuapapal cat 1 " (= quapapalcatl of Hernandez, 1648, not mentioned; usually regarded as Corytophanes hemandezi fide Smith, 1969: 6, 13), sourced from "Seba, p. 132, pi. 82, fig. 1;" and (3) Rana marina3 Amerioana3 perhaps from Mexico (locality not indicated), but almost certainly the species commonly regarded as Bufo marinus (alternatively, in Mexico, as Bufo horribilis) , sourced from "Seba, p. 120, pi. 76, fig. 1." 1 75^a This work is a translation into French of the 1743 book. The section on amphibians and reptiles (pp. 47~ 56) contains virtually the same material as the earlier work; the names are mostly in the French vernacular, however — the "generic" names of Testudo3 Rana and Laoerta do not occur. 1754b The greatest of Klein's works is his "Systeme Naturel du Regne Animal" which was put forth as the equivalent of Linnaeus' "Systema Naturae," as suggested by his subtitle for the first of the two volumes: "Containing the Classes of Quadrupeds, Birds, Amphibians, according to the method of M. Klein; with a notice of that of M. Linnaeus for the same animals; and the Order of Fishes according to the arrangement of Artedi." The frontispiece is a stylized portrait of Klein in his imposing study- museum (Fig. 1) . The organization of amphibians and reptiles (as now understood) does not, however, follow the pattern implied by the volume's subtitle; only two orders were recognized: (l) the Reptilia, including turtles, anurans, crocodi 1 ians, salamanders and lizards, and (2) the Serpens, Bulletin Maryland Herpetol og i cal Society Page 177 Volume 20 Number 4 December 1984 including caecilians, amph i sbaen i ds and snakes. Ten genera were recognized in the Order Reptilia: Testudo 3 Rana, Bufos Crocodilus, Lacerta, Salamcmdra , Cordylus, Scincus , Seps and Cameleo3 five with species of probable Mexican origin. However, the frequent early French practice of eschewing Latin names for animals, in favor of vernaculars, was followed more or less consistently throughout; their use appears to have been incidental. The genus Testudo was divided into two groups of species, much as in his 1751 work, but none was attributed specifically to Mexico. The only species mentioned that do occur there are the sea turtles; their accounts are, however, difficult to assign to species, especially since the seven (or eight) accounts are in part just references to figures or descriptions in the literature without assignment to species (e.g., no. 6 states "Seba, p. 127, pi. 79, figs. 4, 5, 6, 7, gives a figure of three small sea turtles.") . Under the genus Rana only R. marina Americana was mentioned, with the same source given in his 1751 work, but with only "American sea" as a local i ty . In BufOj however, two species, ”B. Brasiliensis " and B. B. orbicu- latus" (nos. 2 and 8 respectively) were treated, both stated to be from Brazil, but given the vernaculars of "aquaqua" and "aquaquaquan ," respectively. Those are names derived from Hernandez ( 1 648) , although not so stated. The first was identified by Smith (1959*. 5) as B. marinus3 but the second name was not found in Hernandez1 work and may therefore be a variant created by Klein. They were sourced from "Seba, p. 1 1 4 , pi. 71, figs. 6, 7" and "Seba, p. 116, pi. 73, fig* 1," respectively. Hence the same species, Bufo marinus, was entered under at least two different names, in different genera, in this work - a not surprising error in view of the rather crude drawings Klein attempted to deal with. The third genus, of "lizards," was divided into two sections, one including crocodi 1 ians, the other true lizards. The first section included three American species, all occurring in Mexico although that country was not mentioned. One, referred to as Crocodilus Americanus (Seba, p. 167, pi. 106, fig. 1) is presumably Crocodylus acutusj the "Crocodile des Indes Occ i denta 1 es, " or "Alligator^" sourced from Catesby, is certainly Alligator mississippiensisj and his " Caimans 3" from "Ceylon, Africa and America" (Seba, vo 1 . 1, pis. 103-105, fig. 3) presumably included the genus Caiman as now known. The second division of "lizards" contains the same fourteen species from Mexico, or possibly from there, as noted in his 1751 work in the genera Lacerta, Cordylus and Cameleo3 although Latin names were not used for most of them in the 1754 work. In the Order Serpens, six "genera" are treated, but none is given an explicit name. The kinds of animals recorded in the literature that are assignable to each genus are listed, and names used by the cited authors Page 178 Bulletin Maryland Herpetol og ica 1 Society Volume 20 Number 4 December 1984 are usually given, but one is left in the dark to know what Klein would call them. The first genus consisted of two species of ichthyophiid caecilians, for which other authors used the name Cecilia. The same ten species of the second genus, defined as having grooves around the body, also consisted of caecilians, of families other than the I chthyoph i i dae, and of some amph i sbaen ians, none clearly from Mexico. The third genus consisted of some seven species with small scales on the abdomen, but no grooves around the body. Species referred by others to Cecilia, Amphisbaena and Scytale are included; one of the latter name is listed for “New Spain" (= Mexico), sourced from “Seba, vol . 2, p. 4, pi. 2, figs. 3, 4“ (see no. 23 in the following account of the Tentamen Herpetol og i ae) . The fourth genus consisted of 27 numbered species, as well as some 25 merely listed from Seba. The genus was characterized as having strap¬ like ventral scales, and judging from the names used by other authors included species of the families, as now understood, Colubridae, Elapidae and those Viperidae with large head scales. Genera cited from other authors include Colubev , Naia, Sibon, Ahaetulla, Petola , Malpolon and Vipera. Nine “species," to judge by name or locality citation, were from Mexico: (1) a “cencoatel" (= cencoatl), whose name originated with Hernandez (1648), not mentioned, interpreted by Smith (1969: 5, 6, 7) as three different species ( Spilotes pullatus, Pituophis deppei and perhaps some species of Leptodeira) , but sourced from Linnaeus' Systema Naturae and from Seba, vol. 2, p. 18, pi. 16, figs. 2, 3; (2) an “apachy- koatl" bearing an Aztec name, but not cited in Hernandez; we assume it is a Mexican species, sourced from four different works, including Seba, vol. 2, p. 21, pi. 20, fig. 1; (3) a spectacled snake from New Spain, sourced from Seba, vol. 2, pi. 97, fig. 4; (4) another “cencoalt" (= cencoatl), sourced from Seba, vol. 2, pi. 26, fig. 1; (5) a “tetzauch- coalt" (= tetzauhcoatl ) , stated to be from Brazil, but bearing an Aztec name, hence probably from Mexico; Hernandez (1648) used the name for only one snake, apparently a Geophis (Smith, 1969: 9), although the same name was applied to skinks and Bipes , neither of which could figure in Klein's “Serpens;" sourced from Seba, vol. 2, pi. 77, figs. 2, 3, pi . 79, fig. 12, and pi. 80, fig. 1; (6) a “tlehua," which name Hernandez (1648, not mentioned) applied to a rattlesnake, perhaps Crotalus polystictus (Smith, 1969: 8), but certainly not so intended by Klein, who put rattlesnakes in another (the sixth) genus; sourced from Seba, vol. 2, pi. 59, fig. 1; (7) another “tlehua," from New Spain, sourced from Seba, vol. 2, pi. 84, fig. 1 (assigned to a “Petzcoal" also in the incertae sedis; see no. 7 of that list, following); (8) a “xalxal hua" (= xaxalhua) from Mexico, regarded as Pseustes poecilonotus by Smith (1969: 9), as the name was used by Hernandez (1648) (not mentioned), but sourced from Seba, vol. 2, pi. 77, figs. 4, 5; and (9) a “depone" (= dopone of Hernandez, 1648, not mentioned), from Mexico, regarded as Oxybelis fulgidus by Smith (1969: 11), but sourced from Seba, vol. 2, pi. 61 . Bulletin Maryland Herpetol og i ca 1 Society Page 179 Volume 20 Number 4 December 1984 Klein's fifth genus of "Serpens" was characterized as having large ventral s, small head scales and no rattles. Some fourteen "species" were included, mostly viperids (as now understood), but apparently also the. boids. One snake from Mexico was cited, sourced from Seba, vol . 2, p. 104, pi . 98, fig. 1 . The sixth genus contained the rattlesnakes, of perhaps seven species, among which Hernandez' (1 648) "teuht 1 acet-zauhqu i " (= teuht 1 acozauhqu i) from Mexico was cited. That name was interpreted by Smith (1969:8) as referring to Crotalus basiliscus and C. durissus, No other source was cited by Klein for that species (although in the Tentamen of 1755 this name was sourced from Seba, pi. 95, fig. 2, referred in 1754b to "Serpens crotalophora" of America). Following the accounts of these six genera, Klein listed 91 figures from Seba of snakes he could not place in his "system;" they were incertae sedis, and among them are at least eight from Mexico: (1) the "ataligato" of Mexico, characterized in Hernandez ( 1 648) by "a totally implausible fable" (Smith, 1969:10), sourced from Seba, vol. 2, pi. 77, fig. 6; (2) "chiametla," from "America," sourced from Seba, vol. 2, pi. 61 , fig. 4, which serves as the type for Coluber chiametla Shaw, 1802, a senior synonym of Drymobius margaritiferus (Schlegel, 1837), but suppressed by the International Commission on Zoological Nomenclature in Opinion 1246 (1983) (see also Smith, 1965 and 1967, and Smith and Smith, 1979); (3) an "emperor of Guadalajara," from Mexico, revered for foretelling the future, sourced from Seba, vol. 2, pi. 1, fig. 1; (4) the "macacoatl," from "America," using Hernandez 1 (1648, not mentioned) name which (Smith, 1969: 7) applied to Boa constrictor , but sourced from Seba, vol. 2, pi. 79, fig. 3; (5) a "macoatl," which probably refers to the "second" "macacoatl" of Hernandez (1648, not mentioned), from Mexico, sourced from Seba, vol. 2, pi. 73, fig. 1; (6) a "n i nboo-quanque cholla," from Mexico, sourced from Seba, vol. 2 pi. 77; (7) a "petzcoal" (= petzcoatl) from Mexico, using Hernandez' (1648, not mentioned) name, which Smith (1969:8) regarded as possibly Dermophis mexicanus , but sourced from Seba, vol. 2, pi. 84, fig. 1, and certainly there not a caecilian; and (8) the "tamaca i 1 1 a-hu i 1 ia" from Mexico (= temacu i 1 cahu i 1 ia of Hernandez, 1648, not mentioned, which Smith, 1969:10 regarded as strictly a fable), but sourced from Seba, vol. 2, pi. 98, fig. 1 (the same source that was used for the Mexican species of his 5th genus; see preceding). 1755 The "Tentamen Herpetol og iae" is written entirely in Latin, and is of special interest as the first published, strictly herpetolog i cal synopsis, although Klein's interpretation of the word differs extensively from its current meaning. It is thus a landmark of sorts in the field, yet is quite a rarity: of the over 750 U.S. institutions cooperating in compilation of the National Union Catalog of Pre-1956 Imprints, only six reported having a copy (Amherst, Boston Public, Cornell, Library of Congress, University of California at Berkeley, Yale). Another copy is in the personal library of Kraig Adler, and we possess two. Page 180 Bulletin Maryland Herpetol og i cal Society Volume 20 Number 4 December 1984 Special attention should be directed to the strangest inclusion of the Tentamen: the only figure of a reptile occurring in it: a fantastic, mythical lizard on pi . 1, shown with a cylindrical body, two powerful, 4-toed hind legs, no forelegs, a stout tail little longer than body, a somewhat skinklike head and wel 1 -devel oped eyes (Fig. 2). It is like no known animal. The only reference to it (p. 50) indicates that it was sourced from a book, "Thesaurus animal ium vivis coloribus egregie pictorum," formerly possessed by "Jobus Lodolphus" (= Hiob Ludolf, 1624- 1704), and that he was uncertain of its proper classification. The animal was not described or illustrated in any of Ludolf 's works, all of which listed in the National Union Catalog we have checked; the book referred to in Ludolf 's library was certainly by some other author, and its identify we have not been able to determine. That the animal (mythical, certainly) was an exasperating incertae sedis for a system of classification recognizing only quadrupeds and limbless "herpeta" is readily understandable. Klein placed his "herpeta" (i.e., herpetozoons , as he interpreted them) in two orders: reptiles in the Order Anguis, worms in the Order Vermis, thus differing markedly from his "system" of the previous year. Little space was allotted to the Order Vermis - only pp. 58~72. Only a little more than one page (55“56) was devoted to "amph i sbaen i ds" (which included some caecilians), hence the bulk of the text concerns snakes . The treatment of Vermis is also strange. There is a brief account of three "classes" (given a lower rank than Order) - Lumbricus, Taenia and Hirudo - with the account ending on p. 66, followed by the two plates and an article (pp. 67“ 72) by J. A. Unzer on "Observat io. . . de Taeniis." All figures (8) on pi. 2, and all but one of the figures (4) on pi . 1 represent the anatomy of parasitic worms (nematodes), accompanying Unzer's article (see his obscure references to those figures on p. 70). Klein's "Order" Anguis was credited with two "classes," neither designated un i nomi na 1 1 y . The first class, by far the larger, was described as having a distinct head and tapering tail, the second class an indis¬ tinct head and blunt tail. The latter class contained two genera - Soytale and Amphisbaena , with 17 and 15 species respectively. The first class was regarded as containing three genera: Vipera, with in effect four subgenera (one the equivalent of viperids as now known, excluding rattlesnakes; one for the rattlesnakes; one for elapids as now known; and one for "vipers" lacking enlarged teeth, hence actually a mixture of non- poisonous snakes), totalling 78 species; Coluber , with 165 species arranged in eight groups of seemingly lesser rank - perhaps similar to species-groups - than those of Vipera ; and Anodon , a supposedly edentu¬ lous genus of five species. 1 Bulletin Maryland Herpetol og i ca 1 Society Page 1 8 1 Volume 20 Number 4 December 1984 The account for the first class terminates with a listing of 14 species of snakes described by Catesby and 33 by Linck. These 47 were regarded by Klein as essentially incertae sedis, along with the two-legged monstrous lizard illustrated on pi . 1 . The species accounts are numbered consecutively in each genus or subgenus. They are brief, of but a few lines, but usually include some sort of characterization, often a locality, and always a source citation from the literature - usually Seba but occas ional 1 y some other - and authority. Many of the species accounts have a marginal rubric giving the species name, in much the same style as Linnaeus' Systema Naturae, although nowhere is that work mentioned in the account of members of his Order Anguis, although frequent references occur in the accounts of members of his Order Vermis. Each of the higher categories is provided with a characterization, and sequentially lettered footnotes (from a to Zzz) at the end of each section give additional information widely representative of the literature, Twenty-four species presumably or certainly from Mexico are treated, (l) Vipera depone , p. 9, no. 13, wa s sourced from "Seba, pi . 92, 11 hence is not the same as the "depone" of his 1754(b) work (see Coluber pullatus , no. 13 of the following accounts), nor is it Hernandez' ( 1 648 , not mentioned) "dopone," from which the name was derived, as interpreted by Smith (1969:11), to wit Oxybelis fulgidus. (2) Vipera crotalophora , p. 16, no. 4, was designated "teutlacot- zouphi," also cited in his 1754(b) work, but in the Tentamen the source was specified as "Seba, pi. 95, fig. 2." The name was clearly derived from Hernandez1 ( 1 648 , not mentioned) "teuht 1 acozauhqu i , " identified by both Duges (1889) and Smith (1969) as Crotalus basiliscus or C. durissus. (3) Vipera conspieillaris altera , p. 18, no. 12, from New Spain, sourced from Seba, pi. 97, fig. 4, is the same as the "spectacled snake" listed in the 1754(b) work, mentioned in the preceding account as the 3rd species in the 4th genus. (4) Ecacoatl , p. 17. A footnote referring to an account on p. 16 of the rattlesnake subgenus" ("Caudisona, Americana1) of Vipera describes in considerable detail the Ecacoatl, from several sources, none of them Seba or Hernandez. The latter author is, however, presumably the ultimate source for the name. His ecacoatl was described as a large, 7-striped snake of several colors, interpreted by Smith (1969) as Mastioophis taeniatus . Obviously Klein had as much difficulty distinguishing venomous from non-venomous species as Hernandez did. (5) Vipera rietu aanino , p. 19, no. 3, was named the "tetzauhcoat 1 ' but was. sourced from a single plate in Seba (80, fig. 1) rather than the three cited for the same name in his 1754(b) work. In the latter work the species was cited for Brazil, but in the Tentamen another Brazilian Page 182 Bulletin Maryland Herpetol og i ca 1 Society Volume 20 Number 4 December 1984 species, " V . thalassina," is given the same name (spelled "tet rauchoat 1 , " but sourced from a plate in Seba (96, fig. 2) still different from any of the three cited for the species in his 1754(b) work. See preceding di scussion. (6) Vipera oculea, p. 20, no. 8, was called "tamacuilla huilia," which name was sourced from Hernandez1 ( 1 648 , so specified) "temacuilca- huil ia," interpreted by Smith (1969) as strictly fictitious. As in Klein's 1754(b) work, the species was sourced from Seba, pi. 98, but cited in both the 5th genus and in the incertae sed i s (no. 8 of that list in the preceding account). (7) Vipera divinatrix and irrrperatrix regni Mexicani, p. 20, no. 11, is the same snake referred to in the 1754(b) work as the "emperor of Guadalajara (misspelled Quadalajara in both works). See preceding dis¬ cussion. Interestingly, the names imperator (of Daudin, 1803) and diviniloquax ( diviniloquus Laurenti, 1768) have long been recognized as junior synonyms of Boa constrictor , a subspecies of which, occurring in Mexico, is still known as imperator. (8) Coluber apachykoatl , p. 25, no. 3, is the same as the snake bearing the same specific name in the 1754(b) work. See preceding dis¬ cussion, to which nothing can be added from the Tentamen. (9) Coluber mexicanus , p. 26, no. 13, was sourced from Seba, pi. 30, fig. 1. Its identify is uncertain. (10) Coluber xajxalh.ua, p. 31, no. 56, is identical with the "xaxalhua" of the 1754(b) work; see preceding discussion. (11) Coluber tlehua, p. 31, no. 57, is the same as the second "tlehua" of the 1754(b) work, sourced from Seba, vol . 2, pi. 84, fig. 1 (which was in 1754b also the source for the "petzcoal" of the incertae sedis). The footnote (tt) on p. 32 merely states that Vipera flammea of the literature is not the same. See preceding discussion. (12) Coluber margariticus , p. 33, no. 68, was not included in the 1754(b) work, but although lacking any locality citation, is here included since it apparently is Drymobius margaritiferus (Schlegel, 1837), a species widespread in Mexico. It was sourced from "Seba, pi. 22, fig. 2." (13) Coluber cencoatl, p. 34, no. 70, has the same source as the second "cencoatl" of the 1754(b) work. As noted previously, the name was used by Hernandez (1648) for at least three species; very likely Pituophis deppei is the one to which this account refers. ( 1 4) Coluber chiametla , p. 38, no. 121, is the same as the "chiametla" of the 1754(b) work; see preceding discussion. Bulletin Maryland Herpetol og i ca 1 Society Page 1 83 Volume 20 Number 4 December 1984 (15) Coluber pullatus y p. 38, no. 122, is the "depone" of the 175Mb) work, being based on Seba's vol . 2, pi. 61 , fig. 2. The name presumably pertains to the species now known as Spilotes pullatus. (16) Coluber petlaeoatl , p. 38, no. 123, sourced from Seba, pi. 63, fig. 1, was not mentioned under that name in Hernandez (1648), nor in the 1754(b) work, although a "petzcoal" was listed there (see preceding discussion), sourced from Seba, vol. 2, pi. 84, fig. 1, which in the Tentamen was assigned to Coluber tlehua (no. 11 above), and in the 1754(b) work to the second tlehua (no. 7 of that account, under the 4th genus) . (17) Coluber ataligato , p. 40, no. 1 38 , is the same as the "atal igato" of the 1754(b) work (see discussion above of the incertae sedis, no. 1, of that work). (18) Coluber maeacoatl , p. 40, no. 139, is the same as the "macacoatl" of the 1754(b) work (see discussion above of the incertae sedis, no. 4, of that work). 09) Coluber capitali fascia latiore, p. 40, no. 139, sourced from Seba, pi. 80, fig. 2, from "New Spain," may be Coniophanes imperialis . The 1754(b) work lists this figure as the source for an incertae sedis, "Bayhapua," an African snake. (20) Coluber petzcoatl , p. 40, no. 1 43 , sourced from Seba, pi. 84, fig. 2, is not the same as the "petzcoal" of the 1754(b) work (Seba, pi. 84, fig. 1). The Tentamen species is arboreal, perhaps Oxybelis aeneus . (21) Coluber cencoatly p. 41 , no. 1 58 , was perhaps intended to be the same as the first "cencoatl" of the 1754(b) work (see no. 1 of the 4th genus in account for that work), but it actually is sourced from Seba, pi. 16, fig. 1, rather than from figs. 2 and 3. (22) Coluber tetzauhaoatl , p. 42, no. 165, was sourced from Seba, pl. 77, figs. 2, 3, one of three sources given in the 1754(b) work for the "tetzauhcoat 1 " of the 4th genus (see no. 5 under that genus in preceding account). Why this snake, bearing a name derived from Hernandez ( 1 648 , not mentioned), was stated to be from Brazil is uncertain. (23) Soytale ex nova Hispania 3 p. 53, no. 5, sourced from Seba, pl. 2, figs. 3, 4, is of uncertain identity. (24) Scytale nixboa quanquecholla3 p. 54, no. 1 4 , sourced from Seba, pl . 77, fig. 1, is the same as the "n i nboo-quanque cholla" of the incertae sedis (no. 6 in the preceding account) of the 1754(b) work. Hernandez1 (1648) "nexoa" was undoubtedl y the source of Klein's names "nixboa" and"ninboo." Smith (1969: 10) interpreted Hernandez' species as Masticophis flagellum3 known to occur in the vicinity of Huauquechula (whence presumably Klein's "Quanquechol 1 a") , Puebla. Page 1 8 4 Bulletin Maryland Herpetol og i ca 1 Society Volume 20 Number 4 December 1984 Numerous discrepancies exist between the 1754(b) and 1755 works, in the areas of overlap, as might well be expected since the latter is much more detailed. A number of differences have been noted in the preceding discussion, but others should be noted. Four "species" of the 1754(b) work are not dealt with in the later review, and seven of the latter are not noted in the former. Omitted in 1755 are: (l) an allocation of the Seba, pi. 16, fig. 3, one of the two sources for the 1754(b) "cencoatel" (4th genus, 1st species of account herewith); two Coluber eeneoatl species were included in the Tentamen, but neither sourced from Seba, pi. 1 6 , figs. 2 and 3 that were the basis for the 1754(b) "cencoatel;" the fig. 3 of pi. 16 in Seba was in 1755 used as the source for Coluber eoyuta of Brazil; (2) the "tlehua" based on Seba, pi. 59, fig* 1 (the Coluber tlehua of 1755 was sourced differently); (3) the "macoatl" based on Seba, pi. 73, fig. 1; and (4) an allocation of Seba, pi. 79, fig* 12, which in 1754(b) served as one of the three sources for the second "cencoalt" (the other two were allocated to Vipera rietu eanino and Coluber tetzauhcoatl) . The species of the Tentamen not represented in the System are (1) Vipera depone of Seba, pi. 92 (the 1754(b) "depone" has a different source); (2) Coluber petlaeoatl of Seba, pi. 63, fig* 1 (the 1754(b) "petzcoal" has a different source, allocated in 1755 to Coluber tlehua and in 1754(b) used as the source also for the second "tlehua") \ (3) Coluber oapitali fascia latiore of Seba, pi. 80, fig. 2; (4) Coluber petzeoatl of Seba, pi. 84, fig. 2 (that figure served in the 1754(b) work as the source for an African snake, "bayhapua;" the "petzcoal" of 1754(b) was based on Seba, pi. 84, fig. 1, which served as source also for the second "tlehua" of that work, and for Coluber tlehua of 1755; (5) Coluber eeneoatl of Seba, pi. 16, fig. 1 (two "eeneoatl" species were included in the 1754(b) work, but neither was based on this figure); (6) Coluber margaritieus of Seba, pi. 22, fig. 2; and (7) Coluber mexiaanus of Seba, pi. 30, fig. 1. 1760a — * This book is a translation into German of the 1751 volume, hence with the same content. The herpetol og i cal pages extend from 285 to 381. 1760b This work likewise is a translation into German of the 1751 Latin discourse, and is not binomial in its nomenclature. A few annotations not in the original Latin have been added, but the organization and taxa are the same. The herpetol og i cal pages extend from 105 to 131. Bulletin Maryland Herpetol og i ca 1 Society Page 1 85 Volume 20 Number 4 December 1384 Summa ry The seven works by Jacob Theodore Klein that pertain to herpetology have no taxonomic importance, most of them being prelinnaean and the two postlinnaean works being non-binomial and therefore not acceptable, according to the International Code of Zoological Nomenclature, for use in nomenclature. The herpetol og i ca 1 accounts were drawn from the litera¬ ture, with greatest reliance upon volumes one and two of Seba (1734-1735), but including also early editions of Linnaeus' Systema Naturae and several other prelinnaean works. His survey of the literature was thorough, and his classification at higher category levels was to a considerable extent original and unique. It was however handicapped by predication on superficial similarities gleaned from descriptions, without the benefit of insights resulting from study of the animals themselves. His analyses did however lead to creation of the word "herpetology,1 although with a distorted understanding that combined snakes, legless lizards, amph i sbaen i ds, caecilians and worms as related groups on the superficial bases of common possession of an elongate body, absence of limbs, and movement by sinuous undulation. Herpetozoons were thus thought of by Klein as the counterpart of quadrupeds, among which he recognized turtles, anurans, salamanders, lizards and mammals. Klein's "system" of classification was, therefore, spurious - a blind alley in the evolution of understanding of the animal kingdom. Having been devised, however, it served to direct the efforts of others into more fruitful channels. Considerable variation in higher-category terminology existed in comparison of early and late works, and even greater confusion is evident in the lower categories. An examination in some detail of the 45-50 "species" noted for Mexico reveals very nebulous concepts tied to often crude or imaginary descriptions and figures in the literature. Klein's works added nothing to an understanding of the Mexican herpetofauna , and indeed clouded the picture. No reason exists to think that the treatment of other herpetofaunae was any better. Hence Klein's herpetol og i cal contributions remain strictly of historical interest, even though his obviously prodigious struggle to discern realities of classification deserve admiration. At least herpetologists of today and the future properly should be conscious of the fact that Jacob Theodore Klein was the "father" of the name for their discipline, and they can be grateful for that detailed unique illustration of the bizarre, mythical, "duobus pedibus lacertinis." Acknowledgments We are greatly indebted to Dr. Kraig Adler for counsel and for xeroxes of the herpetol og i cal sections of Klein's works of 1751 and 1754. Dr. Michael Preston k i ndl y hel ped with translation of Latin passages of critical importance. Page 1 86 Bulletin Maryland Herpetolog ical Society Volume 20 Number 4 December 1984 SYSTEME NATUREL D U REGNE ANIMAL , P A 1 CLASSES , FAMILIES OU ORDRES, CEN’RES ET ESrECCS. A V E C USE NOTICE DE TOUS LES ANIMAUXi Les noms Grccs , Larin j, Ac vul^aircs , x i o i. A PARIS, Quay des A ugufUns , C Volume 20 Number 4 December 1984 Klein, J.T. 1760a. Klassif ication und kurze Geschichte der vierfussigen Thiere, aus dem Lateinischen iibersetzt, und mit Zusatzen vermehret, nebst einer Vorrede von Friedrich Daniel Behn ... Lubeck, Schmidt. 381 pp., 5 pis. (none herpetol og i cal ) . Naturlich Ordnung und vermehrte Historie der v ierfuss i ngen Thiere. Danzig, Schuster. 8 1 , 1, 144, 12 pp. Locupl et i ss imi rerum natural ium thesauri accurato descrip- tio, et iconibus artif iciosissimus expressio, per universam physices h i stor i am. . . . Amsterdam, Jansson i o-Waesberg ios and J. Wetstenium and J. Smith. 4 vols. (vol. 1 appeared in 1734, vol. 2 in 1735, vol. 3 in 1758, and vol. 4 in 1765. All of Klein's references to this work must have pertained to the first two volumes, since he died in 1759 and is unlikely to have incorporated anything from vol. 3 in his writings.) Smith, Hobart M. 1965. Coluber chiametla Shaw , 1 802 (Reptilia, Serpentes) : pro¬ posed rejection as a nomen obi i turn. Bull. Zool . Norn., 22(4) : 235-236, pi . 5 • T96T. Additi onal comment on the proposed rejection of Coluber chiccmetla Shaw, 1 802 , Bull. Zool. Norn., 24(5) :269. 1 969 . The first herpetology of Mexico. Herpetology 3(1) :1-16. _ and Rozella B. Smith 1979* Coluber chiametla Shaw, 1802: revived proposal for suppression under the plenary powers. Bull. Zool. Norn., 35(3) -.184-185. T760b. Seba, Albertus 1734- 1765 — John Johnson, R.F.D. 2, North Bennington 3 Vermont j Malvin L. Skaroff, 1548 Pratt Street 3 Philadelphia 3 Pennsylvania 19124j Hobart M. Smith and Rozella B. Smith, Department of Environmental 3 Population and Organismic Biology 3 and (RBS) , Center for Computer Research in vhe Humanities 3 University of Colorado 3 Boulder 3 Colorado 80309. Bulletin Maryland Herpetol og i ca 1 Society Page 1 89 Volume 20 Number 4 December 1984 NEWS AND NOTES: Office of Fellowships and Grants Smith$onian Institution L "Enfant Plaza. Suite 3300 Washington, D.C. 20560 GENERAL FELLOWSHIP INFORMATION 202 287-3271 3321 The Smithsonian Institution offers fellowships in residence to support independent research and study in fields which are actively pursued by the various bureaus of the Institution. Individuals are selected competitively and are appointed to work under the guidance of professional staff members and to use the collections and facilities of the Smithsonian. The Institution does not generally support work to be done at other insti¬ tutions. It does not offer courses nor does it award degrees. The Smithsonian does not discriminate on grounds of race, color, sex, religion, national origin, age or condition of handicap of any applicant. Six to twelve month pre- and postdoctoral fellowship appointments and ten week graduate student appointments are awarded. Proposals for research in the following areas may be submitted: American Material and Folk Culture American Social, Political, and Military History Anthropology .Linguistics, Archaeology Earth Sciences and Paleobiology Ecological, Behavioral and Environmental Studies - Tropical and Temperate Zones Evolutionary and Systematic Biology History of African Art and Culture History of American, Oriental, and Modern Art History of Design and Decorative Arts History of Music and Musical Instru¬ ments History of Science and Technology Materials Analysis and Conservation of Museum Objects Radiation Biology An applicant must offer a specific and detailed research proposal and must indicate clearly why the Smithsonian is an especially appropriate place to conduct the work proposed. The primary objective of the fellowships is to further the research training of scholars and scientists in the early stages of their professional careers / Predoctoral fellowships are offered to students who have completed preliminary course work and exami¬ nations and are researching the dissertation. Postdoctoral fellowships are generally offered to scholars who have recently completed the doctoral degree. In addition, for 1984-1985 , a few awards will be available for more senior postdoctoral applicants. Candi¬ dates without the Ph.D. but with the equivalent in experience, accomplishment and training may be considered. Graduate student applicants must be enrolled in a program of graduate study and have completed a minimum of one academic semester at the time the appointment begins. Stipends supporting awards are: $18,000 per year plus allowances for postdoctoral fellows; $11,000 per year plus allowances for predoctoral fellows; and $2,000 for graduate students for the ten week period of appointment. Stipends and allowances for pre- and postdoctoral awards are prorated on a monthly basis for periods of less than one year. Individuals .interested in astrophysics or geophysical research should write to: Smithsonian Astrophysical Observatory 60 Garden Street Cambridge, MA 02138 For applications and more information about all Smithsonian fellowships, including the publication, Smithsonian Opportunities for Research and Study, which describes the Institution's bureaus and facilities and lists the professional staff and their research interests, please write to the Office of Fellowships and Grants at the address shown at the top of this sheet. APPLICATION DEADLINE IS JANUARY 15 EACH YEAR Page 190 Bulletin Maryland Herpetological Society Volume 20 Number 4 December 1984 NEW BOOK NEWS: Title: PHYLLIS, PHALLUS, GENGHIS COHEN, & OTHER CREATURES I HAVE KNOWN Author: Fredric L. Frye, DVM Publisher: American Veterinary Publications, PO Drawer KK, Santa Barbara CA 93102 Price: $8.95, softbound (postage paid) LC: 84-70442 ISBN: 0-939674-02-5 Dimensions: 6" X 9" Publication Date: May 1984 Phyllis, Phallus, Genghis Cohen, & Other Creatures I Have Known is a humorous account of the author's 10 years in veterinary practice in Berkeley in the 1960's and '70's. Dr. Frye's specialty is reptiles (a herpetologist is not someone with herpes, he is quick to point out), and this led to some memorable encounters! With the story-telling charm of an urban Herriot, Dr. Frye recounts his adventures with "Playtex the Living Bra" (a snake belonging to a topless dan¬ cer) , "Feather" boa, "Julius Squeezer" (another boa constrictor) and a host of other creatures who, with their equally unusual owners, crossed his thresh¬ old at the Berkeley Dog & Cat Hospital. With Dr. Frye as witty commentator, the reader has a ringside seat as he operates on an alligator in the men's room of a zoo, autopsies a whale, re¬ pairs a snail with a fractured shell, and is the dubious recipient of 90 dozen (1080!) "free" eggs, donated to his veterinary hospital. In these lighthearted episodes, and in the few sobering ones, too, the author comes across as energetic, articulate, and compassionate. Dr. Fredric Frye is someone whom every reader will enjoy getting to know. A sprinkling of masterful cartoons by a fellow veterinarian, Robert M. Miller, enliven these tales. Order directly from American Veterinary Publications, PO Drawer KK, Santa Barbara, CA 93013. $8.95, postage paid. For further information, or to arrange an interview with Fredric L. Frye, contact Roxanne Lapidus, American Veterinary Publications. 800-235-6947. (Calif.: 805-963-6561) AMERICAN VETERINARY PUBLICATIONS, INC. DRAWER KK. SANTA BARBARA. CALIFORNIA 9310? Bulletin Maryland Herpetol og i ca 1 Society Page 191 Volume 20 Number 4 December 1984 NEW BOOK NEWS: NOW PUBLISHED! THE TURTLES OF VENEZUELA by Peter C. H. Pritchard and Pedro Trebbau This book is the first in-depth treatment of a major South American turtle fauna. It covers all turtles known from Venezuela including the matamata and other sidenecks (11 species), tortoises (2 species), pond and land turtles (5 species and subspecies), and the sea turtles (5 species), together comprising half of the turtle species described from the South American continent. There is an extensive discussion of the distribution and zoogeography of South American turtles and a key to species (in both English and Spanish). Each species account consists of a synonymy followed by a diagnosis; a detailed description (including shell, soft parts, color, and sexual dimorph¬ ism); and sections on size and growth, distribution, geographic variation, habitat, feeding, reproduction, economic importance, and vernacular names. The family accounts give a detailed review of the fossil history and present distribution of all genera, worldwide, but with emphasis on South America. There is also a comprehensive bibliography and a list of locality records from throughout the entire continent for all Venezuelan taxa. The book is beautifully illustrated. There are 48 full-page plates in color, 26 of which are original watercolors and the remainder a collection of 165 photo¬ graphs of both turtles and their habitats. In addition, there are two distribu¬ tion maps for each species: a spot map showing the detailed Venezuelan distribution and another map showing the continent-wide range. The book is 414 pages, 8% x 11 inches (21.5 x 28 cm), bound in buckram, price US $45. A special leatherbound patron’s edition, in two volumes, is US $300. A four-page ad with sample color plates was published in the December 1982 issue of Herpetological Review , and copies may be obtained on request from Dr. Douglas H. Taylor, Department of Zoology, Miami University, Oxford, Ohio 45056, USA. Orders may be placed with Dr. Taylor. Please make checks payable to “SSAR.” All USA orders are postpaid; shipments outside the USA will be charged only the additional shipping costs in excess of domestic rates. Overseas customers must pay in USA funds or by International Money Order, or may charge to MasterCard or VISA (give account number and expiration date). A complete list of Society publications and membership information can be obtained from Dr. Taylor. The Society publishes Journal of Herpetology, Facsimile Reprints in Herpetology, Herpetological Review, Herpetological Circulars, Catalogue of American Amphibians and Reptiles, Contributions to Herpetology, and Recent Herpetological Literature. SOCIETY FOR THE STUDY OF AMPHIBIANS AND REPTILES Page 192 Bulletin Maryland Herpetological Society Volume 20 Number 4 December 1984 NEW BOOK NEWS: A New Reprint HERPETOLOGY OF ARABIA by John Anderson with an extensive introduction by Alan E. Leviton and Michele L. Adrich including a new checklist of Arabian amphibians and reptiles In 1896 John Anderson published the book A Contribution to the Herpetology of Arabia , with a Preliminary List of the Reptiles and Batrachians of Egypt, a pioneering effort to summarize the herpetology of those regions and, to this day, the only herpetology of the Arabian Peninsula. The original book is exceedingly rare, probably published in an edition of no more than 100 copies, and is much less known than Anderson’s major work on amphibians and reptiles in the Zoology of Egypt series. The Arabian book includes a description of the physical features, a review of the amphibians and reptiles of the Arabian Peninsula including Yemen, an exhaustive bibliography of the herpetology of Arabia, and a checklist of species both of Arabia and of Egypt including the Sinai. This reprint includes a new introduction with a biography and portrait of Anderson, a list of his publications and an up-to-date checklist of the herpetofauna of Arabia, with map. The book is 160 pages, 6x9 inches (15.5 x 23 cm), and bound in buckram; there is one plate in full color. TO ORDER SSAR members, if ordered before 30 November 1984 . US $18 Institutions; Non-members; All orders after 30 November . US $24 SSAR members must place their orders now to take advantage of the special pre-publication price. The book will be published in December 1984. Send orders to Dr. ^Douglas H. Taylor, Department of Zoology, Miami University, Oxford, Ohio 45056, USA. Please make checks payable to “SSAR”. All USA orders are postpaid; shipments outside the USA will be charged only the additional shipping costs in excess of domestic rates. Overseas customers must pay in USA funds or by International Money Order, or may charge to MasterCard or VISA (give account number and expiration date). A complete list of Society publications and membership information can be obtained from Dr. Taylor. The Society publishes Journal of Herpetology, Facsimile Reprints in Herpetology, Herpetological Review, Herpetological Circulars, Catalogue of American Amphibians and Reptiles, Contributions to Herpetology, and Recent Herpetological Literature. SOCIETY FOR THE STUDY OF AMPHIBIANS AND REPTILES Bulletin Maryland Herpetological Society Page 193 Volume 20 Number 4 December 1984 NEWS AND NOTES: LITERATURE: Fauna Classifieds; 1984 Revisions of U.S. Animal List, Private Collectors Exchange List, U.S. and Foreign Herp List. Free information and sample. Write Bill Gillingham, P 6769, Modesto, CA 95355-0769, (209) 577-6520. Dealer Soc i ety 0 . Box Page 194 Bulletin Maryland Herpetolog ical Society t fi * 1 c Society Publications Back issues of the Bulletin of the Maryland Herpetological Society, where available, may be obtained by wri ting the Executive Editor. A list of available issues will be sent upon request. Individual numbers in stock are $2.00 each, un¬ less otherwise noted. The Society al so publ ishes a Newsletter on a somewhat irregular basis. These are distributed to the membershi p free of charge. Also published are Maryland Herpetofauna Leaf lets and these are available at $. 25/page. information for Authors All correspondence should be addressed to the Executive Editor. Manuscripts being sub¬ mitted for publication should be typewritten (double spaced) on good quality 8i x 1 1 inch paper, with adequate margins. Submit original and first carbon, retaining the second carbon. Indicate where illustrations or photographs are to appear in text. Cite all literature used at end in alphabetical order by author. Major papers are those over 5 pages (double spaced, elite type)andmust i ncl ude an abstract . The authors name should be centered under the title, and the address is to fol low the L i terature Cited. Minor papers are those papers with fewer than 5 pages. Author's name is to be placed at end of paper (see recent issue). For add i t iona 1 information see Style Manual for Biological Journals ( 1 964 ) , American Insti tute of Biological Sciences, 3900 Wi scons in Avenue, N.W. , Washington, D.C. 20016. Price is $6.00. Reprints are available at $.03 a page and shoul d be ordered when manuscripts are submitted or when proofs are returned. Minimum order is 100 reprints. Either edited manuscri pt or proof will be returned to author for approval or correct ion. The author will be res pons ible for al 1 corrections to proof, and must return proof preferably wi thin 7 days. The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc . 2643 North Charles Street Baltimore, Maryland 21218