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B RARI ES SMITHSONIAN INSTITUT = Sa <= % = ig < KBONG> = We < ~ = KE Tops z * ¢ 4 £& \eeae § = yD = S »\ Un, = ‘ a NN \sy a a : 7 a wr] P| - 9 - “a ais i 7 Tras a aa THREE NEW SPECIES OF GEKKO AND REMARKS ON GEKKO HOKOUENSIS (LACERTIFORMES, GEKKONIDAE) Kai-ya Zhoul, Yue-zhen Liu!, and De-jun Li2 1 Department of Biology, Nanjing Normal College 2 Department of Biology, Zunyi Medical College In: Acta Zootaxonomica Sinica 7(4): pp.438-446+pls.1-2, published in October 1982. Translated by Hidetoshi Ota3, Tsutomu Hikida3 & Kuang-yang Lue4 3 Department of Zoology, Kyoto University 4 Department of Biology, National Taiwan Normal University SMITHSONIAN HERPETOLOGICAL INFORMATION SERVICE No. 77 1989 SMITHSONIAN HERPETOLOGICAL INFORMATION SERVICE The SHIS series publishes and distributes translations, bibliographies, indices, and similar items judged useful to individuals interested in the biology of amphibians and_ reptiles, but unlikely to be published in the normal technical journals. Single copies are distributed free to interested individuals. Libraries, herpetological associations, and research laboratories are invited to exchange their publications with us. We wish to encourage individuals to share their bibliographies, translations, etc. with other herpetologists through the SHIS series. If you have such items please contact George Zug for instructions. Contributors receive 50 free copies. Please address all requests for copies and inquiries to George Zug, Division of Amphibians and Reptiles, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Please include a self-addressed mailing label with requests. TRANSLATORS' NOTES In preparing the English version from the original (in Chinese, with English summary), we at- tempted to make as literal a translation as possible. However, a few minor changes were made with footnotes (* — ****); these footnotes follow the references. Locality names were written in Conti- nental spellings, followed by Taiwanese spellings in parentheses at their first appearance. We thank R.I. Crombie and G.R. Zug for their assistance and encouragement during the process of preparation of the present manuscript. INTRODUCTION Among the gekkonid genera occurring in China, Gekko is the largest group with the widest range of distribution. Six species and two subspecies have hitherto been known for the genus from China (Stejneger, 1932; Chen, 1969). During 1975 to 1980, Department of Biology, Nanjing Normal College collected 1637 specimens of Gekko from Hebei (Hopei), Shanxi (Shansi), Shaanxi (Shensi), Shandong (Shantung), Henan (Honan), Jiangsu (Kiangsu), Anhui (Anhwei), Hubei (Hupeh), Sichuan (Szuchuan), Zhejiang (Chekiang), Fujian (Fukien), Jiangxi (Kiangsi), Hunan (Hunan), Guizhou (Kweichow), Yunnan (Yunnan), Guangdong (Canton), Hainan (Hainan)*, and Guangxi (Kwangsi) Provinces. Several specimens were also collected from Guizhou Province by Department of Biology, Zunyi Medical College. While studying these speci- mens, three new species were discovered. On the other hand, Gekko japonicus hokouensis from Yanshan (Chainshan) Prefecture, Jiangxi Province, proved to represent a good species. Conse- quently, ten Gekko species** are presently recognized from China as follows: Key to species of Gekko in China Ihe ROStEHL SCUATALCR ILOMT MOSHE «5. 552253 cia caciss vecmscicicen. eodacedieaue censntelvcalamaubisbiseie ciate ass ev 2 Rostralanicontact with(nostril .c..08.seces ee ee BET ES. Be. See 3 2. Body relatively large, longer than 200 mm in total length; tubercles not particularly concen- trated in upper-marein Ofear Opening 3.000.252. 9-2EA Tae rte eed oO ee eet ae G. gecko Body relatively small, shorter than 150 mm in total length; upper margin of ear opening with cluster of enlarged conical tubercles concentrated in high density ....G. auriverrucosus sp. nov. Soe agle with 24 femoralpOres 1M CACI SIGE oc..osiccccasecoucensaiarescscs sot tess sessaancaat G. kikuchii Male with preanal, or preanal-femoral! pores’ (iii ..2i00) oes SR A RA. 4 At psingie enlarged'spur on/each! side of base of tail 1.0050 27. fas. ee ae eee eas on bee on ctee ee 5 Enlarged spurs, two to three, with slight variation, on each side of base of tail .................. 8 SMES DCEWCER COIS CVIGEM (et hen .cieth ck tae nas «Ras eeaed be cots oct oddace ee ee a caee ee aon e tes Re es 6 Webs. between'dreits very Slent or absent <2 AGA A QO CEE, 2A 7 6. Tubercles absent on dorsum of body; male with 7-11 preanal pores ............. G. subpalmatus Tubercles present on dorsum of body; male with 17-27 preanal pores............... G. chinensis 7. Supranasals in contact; dorsal tubercles flat; head and body length reaching 80 mm .............. a stains Mes y cepasts ania coir s ALWin a Sjessidic 5 ae ee ais onadeaNoosasitane sic ace dese tase test G. liboensis sp. nov. Supranasals ‘3. arated by a small scale; dorsal tubercles relatively convex; head and body lenet notpredter faan-7 O WAMN 6.0 b Fig. 1. Gekko auriverrucosus sp. nov. a. Dorsal view of head; b. Ventral view of head. Gekko auriverrucosus Zhou et Liu sp. nov. (Plate I: 2, Fig. 1) Holotype—Male (NNC 80275), collected from Hejin (Hojin) Prefecture, Shanxi Province (alt. 459 m), on 19 August 1980. Allotype: female (NNC 80243), collection date and locality as for holotype. Paratypes: 33 males and 41 females, collected from Hejin, Yongji (Engtsi), and Linyi (Linyi) Prefecture, Shanxi Province. Collector: Xin-rong Xu. Type specimens are deposited in Department of Biology, Nanjing Normal University. Diagnosis—Rostral separated from nostril; upper margin of ear opening with cluster of enlarged conical tubercles gathering in high density; tubercles uniformly scattered in temporal and occipital regions, neck, and dorsal surfaces of body, base of tail and limbs; male with 8-11 preanal pores. Description—Snout about twice as long as eye diameter, distinctly longer than distance between eye and ear opening; diameter of ear opening 0.9-1.5 mm, about 30-44% of eye diameter; rostral twice as broad as high, angulated at midpoint dorsally, separated from nostril; nostril surrounded by first supralabial, supranasal, and two small scales; supranasals moderately enlarged, slightly longer than broad, separated from each other by a minute scale, or in contact with each other medi- ally; supralabials 9-11; infralabials 9-11; mental pentagonal; chin shields forming several rows of transverse arches; first row normally comprising five shields, each slightly longer than broad, median three largest; scales following chin shields and reaching gular region uniform, granular (Fig. 1). About 12 scales between nostril and eye; interorbital scales about 25; upper margin of ear open- ing with tubercle cluster comprising about six enlarged conical tubercles gathering in high density; around jaw angle and preotic region also with enlarged conical tubercles; tubercles uniformly scat- tered among dorsal granular scales, from temporal and occipital regions to base of tail, in 16-20 irregular rows at midbody; dorsal surfaces of forelimbs covered with small tubercles; on dorsal surfaces of hindlimbs, tubercles scattered among granular scales; scales granular in gular region, imbricate in the other part of ventral surface of body; webs between digits rudimentary; underneath dilated portions of toes with lamellae, 6-8 on toe I, 6-8 on toe II, 6-8 on toe III, 6-8 on toes IV, and 7-9 on toe V; male with 8-11, mostly 8-9 preanal pores. Table 1. Measurements (in mm) of specimens of Gekko auriverrucosus. Diameter Axilla Fore- Hind- Total Eye of ear Snout Head groin limb limb Specimens length diameter opening length length length length length Holotype (NNC 80275) 125.5 (62+63.5) 3.2 1.2 6.7 15 28 18.5 24.5 Allotype (NNC 80243) 135.5 (65.5+70) 3.3 13 i/RS) 16 30.5 19 27 Paratypes 11 males 119 (59+60)- 3- 1- 6.3- 14- 25- 16- 24- from Hejin 130 (63+67) 335 ja 7 16 29.5 19 25 16 females 117 (56+61)- 3- 1- 6.4- 14- 27- 16.5- 23- from Hejin 133 (65+68)*** 3.6 | igs ype) 17 33 20 28.5 Tail slightly compressed, with two or three enlarged spurs in each side at base; dorsum of tail covered with tubercles of various sizes; annular grooves in about every sixth to eighth tubercle; venter of tail with a longitudinal row of laterally elongated shields. Dorsal ground color of preserved specimen pale gray; a brown bar from nostril through eye and ear to shoulder; top of head with brown markings; dorsal surfaces of neck and body with 5-6 transverse brown bands; dorsum of tail with 9-13 transverse brown bands; posterior edge of transverse bands in body and tail darkly edged; dorsal surfaces of four limbs also with transverse brown bands; venter of body light reddish yellow. This new species might be easily misidentified as G. japonicus. However, the latter species has a rostral entering the nostril, and lacks a cluster of tubercles in the upper margin of the ear opening. Thus, G. japonicus is actually distinct from the present new species. In the natural habitat, the density of G. auriverrucosus is very high. It prefers to perch on high portions of walls, and occasionally appears on artificially lighted areas to search for prey. In June and July, the present species has its reproductive season. Juveniles collected between 19 and 22 August had already reached 31-32.5 mm in head and body length. All adult females collected on the same date from the same locality with the above juveniles possessed no mature eggs. About 1/6 of the total sample had parasitic mites, especially in high density on digits. Gekko liboensis Zhou et Li sp. nov. (Plate II: 1, Fig. 2) Holotype—Female (TMC 791669), Chengguan (Chengkwang), Libo (Libo) Prefecture, Guizhou Province (alt. 430 m), on 5 July 1979, by Zhi-lu Zhao. This specimen is deposited in Department of Biology, Zunyi Medical College. = b Fig. 2. Gekko liboensis sp. nov. a Dorsal view of head; b. Ventral view of head. Diagnosis—Head and body length exceeding 80 mm; supranasals large, in contact with each other medially; flat, cycloid tubercles among dorsal granular scales, forming about 10 irregular longitudinal rows; webs evident between fingers I, II and III, very slight between fingers III, IV and V. Description—Holotype very large, head and body length nearly 85 mm; snout 1.8 times as long as eye diameter, distinctly longer than distance between eye and ear opening; diameter of ear opening 2 mm, about 40% of eye diameter; rostral broader than deep, upper margin slightly con- caved at midpoint; nostril surrounded by rostral, first supralabial, supranasal, and two small scales; supranasals large, in contact with each other medially; supralabials 12; infralabials 11; mental triangular; median pair of chin shields twice as long as broad, posteriorly entered by a pair of small polygonal chin shields (Fig. 2). About 18 scales between nostril and eye; interorbital scales about 40; flat, cycloid tubercles uni- formly scattered in low density on dorsum, from parietal and occipital regions to base of tail, forming about 10 irregular rows on body; fore- and hindlimbs without tubercles; ventral surface posterior to neck covered with imbricate scales; 10 enlarged scales in preanal region. Underneath dilated portions of fingers covered with lamellae, eight on finger I, eight on finger II, nine on finger III, nine on finger IV, and eight or nine on finger V; rudimentary webs evident Table 2. Measurements (in mm) of a specimen of Gekko liboensis. Diameter Axilla Fore- Hind- Total Eye of ear Snout Head groin limb limb Specimens length diameter opening length length length length length Holotype (TMC 791669) 121.8 (84.8437) 5 2 92 7a\\ 3y/ PISA 365) (regenerated tail) between fingers I, II and III, faintly between fingers III, [V and V; margins of webs attaching to proximal one third of toes; hindlimb much developed, its length 95% of axilla to groin length; underneath dilated portions of toes covered with lamellae, eight on toe I, seven or eight on toe II, eight on toe III, nine on toe IV, and nine on toe V; rudimentary webs evident between toes I, II, III and IV; a single large spur on each side of base of tail; tail regenerated, very short. Dorsal ground color in preservative grayish tan; a brown bar running along lower margin of eye, almost reaching to ear opening posteriorly; dorsal surfaces of neck and body with nine transverse brown bands; dorsal surfaces of limbs also with transverse brown bands; venter of body pale red- dish yellow. This new species greatly resembles G. hokouensis. However, the latter has supranasals sep- arated from each other, and conical dorsal tubercles. Moreover, the head and body length of G. hokouensis is shorter than 70 mm. G. liboensis is rarely observed at Chengguan, Libo Prefecture. Gekko hokouensis Pope Gekko japonicus hokouensis Pope, 1928, Amer. Mus. Novitates 325: 1-2 (Yanshan Prefecture, Jiangxi Province) Pope (1928) regarded this form as a subspecies of G. japonicus, and stated that G. j. hokouensis differs from the nominal subspecies only in the number of cloacal spurs; he noted that the former has a single spur on each side of the base of tail, whereas the latter has two or three spurs. While investigating a large series of specimens, we found that hokouensis has a relatively large spur, measuring about 2.2-3.1 mm for the male and 1.3-2.0 mm for the female in maximum diameter. Although the spur is more or less grooved and incompletely divided in a few males and most females, the outline of the single spur remains apparent in all animals (Plate II: 4-9). On the other hand, japonicus possesses two or three smaller spurs below three larger spurs. The size of each spur is relatively small, and the maximum diameter of the largest spur measured 1.2-1.5 mm in the male and 0.6-1.0 mm in the female (Plate II:10-11). Differences are recognizable between hok- ouensis and japonicus also in the condition of dorsal tubercles as follows. In hokouensis, tubercles are absent on the four limbs, and relatively few around the middle of the body. On the other hand, in japonicus, the dorsal surface of the forearm and shank is covered with tubercles, and the tuber- cles around the middle of the body are in relatively high density (Plate II:2-3, Table 3). Table 3. Comparison of dorsal tubercles in Gekko hokouensis and G. japonicus. Occipital Upper Species N Localities andneck Body am Forearm Thigh Shank Gekko hokouensis 271 17locationsin -/+ - - - - - Six provinces Gekko japonicus 747 50 locations in + ++ - + -/+ + 12 provinces Fig. 3. Distributions of Gekko hokouensis and Gekko japonicus. Ranges of the former and the latter are outlined by broken and complete lines, respectively. Cross-marks indicate localities of specimens of G. hokouensis cited in the present study****. On the basis of the above characteristics, we identified 271 specimens out of 1018 of G. japoni- cus (sensu lato) as hokouensis, and the remaining 747 as japonicus (sensu stricto). The former specimens were collected from 17 prefectures of six provinces, and the latter from 50 prefectures or cities of 12 provinces. The sampling localities of hokouensis are scattered within the range of japonicus (Fig. 3). Conditions of natural habitats also differ between these two gekkonids; while hokouensis has its habitat in montane environments, japonicus is widely distributed in cities and villages of the plain regions. In Jiujiang (Kiukiang) City, for example, japonicus is found in urban area, whereas hok- ouensis in Lushan-haihui (Lushan-haihui), Bailudong (Bailudong) and Guling (Kuling). In Yixing (Ising) Prefecture, japonicus and hokouensis occur in the lowland and montane areas, respectively. The mutual displacement between hokouensis (a mountain dwelling species) and japonicus (a plain dwelling species) around a sympatric area much resembles the displacement between Eremias brenchleyii and E. argus. However, there are some areas where both of these gekkonids are col- lected from the same point at the same time. For example, of the 25 specimens obtained within a city of Chong'an (Chungan) Prefecture, on 26-27 June 1978, 15 specimens were identified as japonicus and the remainder as hokouensis. No intermediate forms were found among the above specimens. Based on the morphological distinctiveness, sympatric occurrence, and ecological differences, we remove hokouensis from the subspecific status of G. japonicus, and regard it as a distinct species Gekko hokouensis Pope. Within China, G. hokouensis is distributed in Yixing and Lishui (Lishui), Jiangsu Province, Tonglu (Tonglu) (Stejneger, 1932), Longquan (Longchuan), Beiyandangshan (Beiyantangshan), Zhejiang Province, Pucheng (Pucheng), Chong'an, Wuyishan (Wuyishan), Shaowu (Shaowu), Youxi (Yousi), Fujian Province, Taiwan Province (Maki, 1923), Jinzhai (Ginshai; Xuzhou Nor- mal University), Huangshan (Huangshan) and Taiping (Taiping; Chengdu Institute of Biology), Anhui Province, Yanshan, Ninggang (Ningkwang), Lushan, Jiangxi Province, Yizhang (Ichang; Zhengdu Institute of Biology), and Jiangyong (Kiangyong)!, Hunan Province. In Taiwan, G. japonicus has been reported from several localities (Chen, 1969). Maki (1923) described speci- mens of “G. japonicus” from Taiwan as possessing a single process in each side of the base of tail, and limbs covered only with granular scales. These characteristics are identical with those of G. hokouensis. Of the specimens of “G. japonicus” collected from Japan, some animals such as the one from Nagasaki (USNM 13563; Stejneger, 1907) were reported to have three spurs in each side at the base of the tail, and tubercles on the dorsum of the shank, whereas others such as specimens from Yamagawa (USNM 31821 and 31822; Stejneger, 1907) and Fukuoka (Okada, 1936: fig. 1) possess a single spur and lack tubercles on limbs. Nakamura and Uéno (1963) noted that G. japonicus has normally a single but occasionally two or three tubercles, and that some animals possess tubercles on limbs. In October 1981, one of us examined specimens under the care of Dr. Shun-Ichi Uéno at National Science Museum, Tokyo. Of the 21 specimens there, 12 from Tokara Is. (0231, 02287, 02293-95), Tokunoshima I. (0924), Yoronjima I. (0524, 02288-90, 02292) and Iriomotejima I. (0475) were identified as G. hokouensis, and the other nine from Tokyo (0236, 02286, 02297-98), Kyoto (02302) and Tsushima I. (0038, 0851, 0853, 02296) as G. japonicus. These results indicate that “G. japonicus” in Japan actually includes both G. hokouensis and G. japonicus (sensu stricto). Gekko scabridus Liu et Zhou sp. nov. (Plate I: 1, Fig. 4) Gekko sp. Hu Shu-chin, Djao Er-mie and Liu Cheng-chao, 1973, Acta Zoologica Sinica 19(2): 155, from Guiyang (Kweiyang) Holotype—Male (NNC 80122), Yongren (Yongzen) Prefecture, Yunnan Province (alt. 1531 m), on 4 Aug. 1980. Allotype: female (NNC 80143), collection date and locality as for holotype. Paratypes: 16 males and 33 females collected from Yongren Prefecture, Yunnan Province, and Miyi (Miyi) Prefecture, Sichuan Province. Collector: Xin-rong Xu. Type specimens are deposited in Department of Biology, Nanjing Normal University. Diagnosis—Tubercles covering dorsal surfaces of body and hindlimbs much enlarged; male with 10-15 preanal pores. Description—Eye relatively large, its diameter longer than half, as long as 51.4-57.1%, of snout length; snout slightly longer than distance between eye and ear opening; diameter of ear opening 0.9-1.5 mm, about 23-41% of eye diameter; rostral rectangular, its breadth less than twice of height; in a few specimens, upper margin of rostral slightly concave dorsally at mid point; nostril surrounded by rostral, first supralabial, supranasal, and two small scales; supranasals moderately enlarged, slightly broader than long, separated from each other normally by a single scale, but in 1 All the locality data without citations of authors or institution are based on the specimens deposited in Department of Biology, Nanjin Normal College. a b Fig. 4. Gekko scabridus sp. nov. a. Dorsal view of head; b. Ventral view of head. some specimens, by two scales, or in contact with each other medially; supralabials 9-11; infral- abials 9-11; mental pentagonal; chin shields longer than broad, median pair largest, outer pair rela- tively small, continuously graded to small granules through three to four rows of small hexagonal scales (Fig. 4). About 12 scales between nostril and eye; interorbital scales about 30; tubercles scattered among dorsal granular scales in high density, from frontal, parietal, temporal and occip- ital regions of head to base of tail, in 17-21 irregular rows around middle of body, those on dor- sum of body extremely enlarged; limbs covered with granular scales dorsally, tubercles present on limbs except for upper arms; tubercles on hindlimbs distinctly enlarged like those in dorsum of body; venter of body covered with imbricate scales; interdigital webbings rudimentary; underneath dilated portions of digits with lamellae, 6-9 on toe I, 6-9 on toe II, 7-9 on toe III, 7-9 on toe IV, and 7-10 on toe V; male with 10-15, mostly 12 or 13, preanal pores. Tail slightly compressed, with two or three enlarged spurs on each side at base; only one spec- imen (NNC 80166) of 51 examined had a single enlarged spur on both sides; dorsum of tail cov- ered with granular scales; annular grooves in about every seventh to ninth row of granules; grooves in proximal one third of tail margined by six to eight enlarged tubercles posteriorly, such tubercles gradually disappearing in remaining portion of tail; venter of tail covered with imbricate scales, median scales enlarged and irregularly arranged, paired or not paired, in distal two-thirds to four-fifths of tail. Dorsal ground color of preserved specimen pale brown; two brown bars from nostril through eye to temporal region; dorsal surfaces of head, body and limbs with irregular brown spots and reticu- lations; 7-9 transverse bars on neck and body; dorsum of tail with 10-14 transverse brown bars; venter of body light reddish yellow. G. scabridus closely resembles G. japonicus. However, these species differ from each other as follows. In G. japonicus, dorsal tubercles on the body and shanks are distinctly smaller than those in G. scabridus, and tubercles are normally lacking on thighs. Moreover, male G. japonicus nor- mally has only 4-8 preanal pores. In the natural habitat, G. scabridus occurs in very high densities, and is observed equally in lighted and dark areas of walls. Of the specimens examined, a few animals possessed parasitic Table 4. Measurements (in mm) of specimens of Gekko scabridus. Diameter Axilla Fore- Hind- Total Eye of ear Snout Head groin limb limb Specimens length diameter opening length length length length length Holotype (NNC 80122) 112.5 (S7.5+55) 3.7 V3 ih 15.7 26 LW (a) 26 Allotype (NNC 80143) 120.5 (58+62.5) 3.8 1.4 7 13507, 27 19 24.5 Paratypes 11 males 114 (57+57)- 3.7- 1.2- 7- 15.4- 25.5-. 17.5- 25- Yongren 138 (64+74) 4.2 1-5 122) 17. 30 20 27 12 females 116.5 (56.5+60)- 3.5- 1.1- 6.3- 14.5- 25- 16.5- 24- Yongren 140.5 (63.5+77) 4 jes 1) i, 30 19 P| mites. Juveniles collected in the beginning of August had reached 28-33 mm in head and body length. All adult females collected in the same date from the same locality with the above juveniles had no mature follicles. REFERENCES Chen, J.T.F. 1969. A Synopsis of the Vertebrates of Taiwan. Commercial Press; Taipei. (in Chi- nese) Hu, S.-C., E.-M. Djao, and C.-C. Liu. 1973. A survey of amphibians and reptiles in Kweichow Province, including a herpetofaunal analysis. Acta Zoologica Sinica 19(2): 149-178. (in Chinese with English summary) Maki, M. 1923. On the gekkonid lizards of Taiwan. Zool. Mag. 35: 193-202. (in Japanese) Nakamura, K., and S.-I. Uéno. 1963. Japanese Reptiles and Amphibians in Colour. Hoikusha; Osaka. (in Japanese) Okada, Y. 1936. Studies on the lizards of Japan. Contribution I. Gekkonidae. Sci. Rep. Tokyo Bunrika Daigaku, B 2(42): 233-289. Pope, C.H. 1928. Four new snakes and a new lizard from South China. Amer. Mus. Novitates 325: 1-4. . 1929. Notes on reptiles from Fukien and other Chinese Provinces. Bull. Amer. Mus. Nat. Hist. 58: 335-487. . 1935. The Reptiles of China. Nat. Hist. Central Asia. Vol. 10. Amer. Mus. Natur. Hist., New York. Stejneger, L. 1907. Herpetology of Japan and adjacent territory. Bull. U.S. Natl. Mus. 58(1): 1- SS: . 1932. The Chinese lizards of the genus Gekko. Proc. U.S. Natl. Mus. 82(3): 1-8. Wermuth, H. 1965. Liste der rezenten Amphibien und Reptilien, Gekkonidae, Pygopodidae, Xantusiidae. Walter de Gruyter & Co.; Berlin. 10 TRANSLATORS' FOOTNOTES * In the original, Hainan Island was included in Guandong Province. However, this island was removed from the latter to form an independent province, Hainan Province, by itself in 1987. ** The original states “10 Gekko species and one subspecies”, but it lists only 10 species in the key. *** ‘Table 1 in the original contains “113 (65+68)”, but this must be a typographical error. **** The figure legend in the original states that the cross-marks in Fig. 3 indicate localities where G. japonicus was collected. But it is evident, from the content of the text, that those marks actually represent sampling localities of G. hokouensis. Plate I. 1. Gekko scabridus Liu et Zhou, sp. nov. Lateral view of head (above), and dorsal view (below). 2. Gekko auriverrucosus Zhou et Liu, sp. nov. Lateral view of head (above), and dorsal view (below). Plate II. 1. Gekko liboensis Zhou et Li, sp. nov. Dorsal view. 2. Gekko hokouensis. Dorsal view of base of tail and hind limbs. 3. Gekko japonicus. Dorsal view of base of tail and hind limbs. 4-9. Gekko hokouensis. Enlarged spurs on right side of base of tail in males (4 and 5) and females (6 to 9). Spurs in 4 and 6 have no shallow furrows, whereas those in the remainders (5, 7, 8, and 9) show more or less developed furrows incompletely dividing the spurs. 10-11. Gekko japonicus. Spurs on right side of base of tail in a male (10) and a female (11). INDEX TO THE BIOGRAPHIES OF HERPETOLOGISTS &@ ICHTHYOLOGISTS COPEIA 1913-1988 MAY 18 1993) CIB RAniE €. J. McCoy, Section of Amphibians and Reptiles Carnegie Museum of Natural History SMITHSONIAN HERPETOLOGICAL INFORMATION SERVICE No. 78 1989 SMITHSONIAN HERPETOLOGICAL INFORMATION SERVICE The SHIS series publishes and distributes translations, bibliographies, indices, and similar items judged useful to individuals interested in the biology of amphibians and_ reptiles, but unlikely to be published in the normal technical journals. Single copies are distributed free to interested individuals. Libraries, herpetological associations, and research laboratories are invited to exchange their publications with us. We wish to encourage individuals to share their bibliographies, translations, etc. with other herpetologists through the SHIS series. If you have such items please contact George Zug for instructions. Contributors receive 50 free copies. Please address all requests for copies’~ and inquiries to George Zug, Division of Amphibians and Reptiles, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Please include a self-addressed mailing label with requests. Let us now praise famous men, and our fathers that begat us. All these were honored in their generations, and were the glory of their times (Ecclesiasticus 44:1). INTRODUCTION The history of a science consists not only of the chronology of development of knowledge and ideas, but equally of the life stories of men and women who contribute to that science. For ichthyology and herpetology a significant part of this human record is contained in seventy-five years of the journal Copeia. Unfortunately this rich source of historical and biographical information has been largely inaccessible, as most of the death notices, obituaries, and biographical sketches in Copeia appeared in the "Editorial Notes and News" section of the journal, and are not included in the journal's annual index. This compilation will provide ready access to biographies in Copeia and, hopefully, stimulate an appreciation for the personalities who fostered modern ichthyology and herpetology. The index consists of three lists: the biographical articles arranged alphabetically by name of subject and keyed to the bibliography by number, an annual list of biographies, and a bibliography. The content of articles is coded as follows: "N" is a death notice only; "B" is a biography; "P" indicates a portrait is included; "L" identifies a biography with a bibliography. The biographies range in length from a few words to several pages. Although the distinction is arbitrary in some cases, the "B" category is reserved for articles that have substantial biographical content. Three major articles in Copeia (Dymond, 1964, (1):2-33; Myers, 1964, (1):34-41; Hubbs, 1964, (1):42-60) are not indexed. Although they contain biographical information, these articles are primarily historical reviews. Much of the credit for documenting the lives of ichthyologists and herpetologists in Copeia belongs to two of our late colleagues, incidentally also my "scientific grandfathers." The first of these, Carl L. Hubbs, had an unusually strong sense of the importance of the historical perspective. Perhaps this was because his career spanned critical years in the transition to the modern era, as did that of his mentor David Starr Jordan. Hubbs was by far the most prolific contributor of biographical articles to Copeia (24 signed articles and dozens of unsigned notes). Moreover, as first ichthyological editor (1930-1937) of the "new series" of Copeia, he pioneered regular incorporation of biographical information in "Editorial Notes and News." George S. Myers is second to Hubbs in number of biographies published in Copeia, but his contributions are models of sympathetic and _ insightful biography. To these two, indeed to all who have contributed to this unique record, we owe our gratitude. I thank Alan E. Leviton for (indirectly) suggesting this project, and M. Therése Giles for enduring patience. BIOGRAPHIES Alcock, A. W.:30 Allen, E. J.:98 Aliten,(W 4B. ¢i3h Allen, William Ray:169 Allis, -JZ.,; Edward Phelps:132 Alm, Gunnar:327 d'Ancona, Umberto:208 Anderson, Jr., James Donald: 419 Anderson, Paul:201 Andersson, Lars Gabriel:149 Andrews, Roy Chapman:188 Angel, Fernand:143, 307 Artedi, Peter:352 Atkinson, N. J.:25 Awerinzew, Sergius:114 Ayers, Howard: 32 Babbitt, Lewis H.:235 Babcock, Harold Lester: 350 Babcock, John Pease: 48 Bacon, Daniel:35 Baranov, Fedor Ilich:373 Barbour, Thomas: 289 Barnard, K. H.:209 Barnhart, Percy S.:157 Basspeed = fant oOhm pe. <9: Bauman, Aden C.:134 Beaufort, Lieven Ferdinand de:264 Bennett, George W.:237 Bensley,’ B. -A.:34 Berezovsky, A. I.:115 Berg, Leo Semenovich:147, 388, 389 Bertin, Léon:172 Bigelow, Henry Bryant: 285 Bishop, Sherman C.:150, 305 Blair, William Franklin: 341 Blanchard, Frank Nelson: 376 Blegvad, Harald:155 Bohlke, James E.:374 Borodin, Nicholas A.:70 Boulenger, E. G.:122 Boulenger, George Albert:66, 395 Bragg, Arthur Norris: 263 Breden, s3ra, C2 9Ma:252 Breyer-de Rooy, P. Je240'5 Brimley, Clement Samuel: 281 Bromund, Fred:108 Brumwell, Malcolm J.:97 Bruun, Anton Frederik:197, 323 Buen, Fernando de:324 Burt, Charles E.:405 Cagle, Fred Ray:407 Cameron, A. T.:135 Camp, Charles L.:417 Carr, irs, “ARCHER. 129i. Chabanaud, Paul:177 Chernov, Sergius Alexandrovich: 203, Clark, Howard Walton:95, 96 Clark, Hubert Lyman:129 Clay, William Marion:277 Clemens, Lucy Wright Smith:55 Clemens, Wilbert A.:210 Cobb, John Nz 257 9380 Cochran, Doris Mable:301 Cockereliy T: Ds *As: 136 Conant, Isabelle:236 Cook, Fannye A.:206 Cooper, Gerald Paul:256 Cope, Edward Drinker:269, 298, 357 Cowles, Raymond Bridgman: 266 Cravtord, ¥Srs; Donald *R. 3195 Creaser, Charles W.:302 Crimmins, Martin Lalor:168 Cunningham, J. T.:42 Curtis, Brian:191 Cyren, Otto:128 Czopek, Juliusz:403 Dahlgren, Ulric:123 Daniel, John Franklin:99 Davies, David H.:329 Davis, =D. Dwight: 221 De VHads 7 «Ce (Oe We 2 141 Ditmars, Raymond L.:375 Donoso-Barros, Roberto: 368 Duncker, Georg:166 Dunn, Emmett Reid:171, 378 Dybowsky, Benedikt:6 Dymond, John Richardson:212, 385 Eager, Grace:137 Eaton, Theodore H.:412 Eddy, Brayton:148 Eddy, Samuel: 414 Ege, Vilhelm:325 Ehrenbaum, Ernst:133 Ehrenreich, Alfred:16 Eigenmann, Rosa Smith:127 348 Einarsson, Hermann: 218 Ellis, Max M.:164 Embody, George Charles:74, 80 Engeling, Gus A.:159 Erwin, Richard P.:21 Evermann, Barton Warren:22, 310 Ewing, H. E.3152 Fassett, Harry C.: 311 Flower, Stanley Smyth:121, 396 Forbes, Stephen Alfred:7 Fowler, Henry Weed:276 Fox, Jr., Wade:282 Frizzell, Don L.:231 Gage, Simon Henry:109 Gaige, Frederick MacMahon:254 Gaige, Helen Thompson: 254 Giltay, Louis Pierre:59 Ginsburg, Isaac:360 Glauert, L.:202 Gloyd, Howard Kay:271 Goff, Carlos Clyde:73 Goin, Coleman J.:273 Goodnoh, Jr., Clifford A.:116 Gordon, Myron:178, 253 Gowanloch, James Nelson:161 Greenbank, John T.:240 Greenberg, Bernard S.:214 Grey, Marion:399 Grinnell, Joseph:82, 88 Grodzifiski, Zygmunt: 402 Gudger, E. W.:173, 364 Ginther, Klaus:290 Haas, Georg: 418 Hamilton, Rodgers D.:145 Hankinson, Thomas L.:45 Hanko, Bela:184 Hansen, Donald F.:238 Harrington, Jr., Robert W.:251 Hart, John L.:386 Hartweg, Norman E.:204 Hass, Robert L.:222 Hay, Oliver Perry:11 Hay, William Perry:126 Henderson, Junius:67 Herald, Earl Stannard: 361 Herre, Albert W.:198 Hildebrand, Samuel Frederick:139, 315, 319, 379,43837 384 Hine, James S.:12 Hollister, Gloria:261 Hoover, Earl E.:72 Hora, Sunder Lal:392 Howard, John K.:213 Hoyt, Southgate Y.:153 Hubbs, Carl Leavitt:342, 353, 365, 391 Hyde, Jesse Earl:52 Ihering, Rodolpho von: 421 Ishikawa, Chiyomatsu: 40 Jacot, Arthur Paul:81 Jenkins, Oliver Peebles:41 Johansen, A. C.:18 Johnson, Ose Eason, 377 Johnson, Richard M.:349 Johnstone, James:26 Jordan, David Starr:15, 292, 296, 387 Jordan, Eric Knight: 306 Juday, Chauncey:106 Kampen, P. N. van:69 Kelly, Howard A.:100 Kendall, William Converse:75, 79 Kishinouye, Kamakichi:8 Klauber, Laurence Monroe: 390 Klugh, A. Brooker:19 Knight, A. P.:46 Koumans, Frederick P.:265 Kuehne, Robert Andrew: 257 Lagler, Karl F.:355 Lahille, Fernando:94 Landreth, Hobart:233 Lantz, Louis A.:366 Lataste, Fernand: 36 Limbaugh, Conrad:322 Lindberg, G. U.:2 Loftus-Hills, Jasper J.:1 Logier, Eugene Bernard Shelley:278 Longley, William Harding: 53 Loénnberg, Einar:320 Lowe, John N.:71 Lubbert, Hans:160 Lutz, Adolpho:93 Lutz, Bertha:345 Lutz, Gualter Adolpho:223 Maki, Moichiro:183 Mann, William M.:189, 194 March, Douglas D. H.:83 Maria, Niceforo:404 Martof, Bernard S.:255 Maskell, F. G.:31 Maslin, T. Paul:393 Matsubara, Kiyomatsu: 331 Mayer, Fritz:179 McGregor, Richard Crittenden: 56 McIlhenny, Edward Avery:268 McMurrich, James Playfair:76, 87 Meade, George P.:288 Merriman, R. Owen: 39 Mertens, Robert: 300 Metzelaar, Jan:4 MichaYowski, Jerzy:219 Miller, Frances Voorhees Hubbs:270 Miller, Richard B.:185 Miller, Richard D.:180 Mieintosihyn iW iC.s320 Miranda Ribeiro, Alipio de:84, 356 Miranda Ribeiro, Paulo de:215 Mitchell, F. J.:224 Mohr, Erna:332 Morgan, Stanley Stewart:111 Mosauer, Walter:57 Mukerji, Dev Dev:60 Miller, Lorenz:313 Myers, George Sprague:272 Nakamura, Hiroshi: 413 Needham, Paul R.:207 Nellemose, (Captain):119 Nelson, Edward William:37 Nichols, John Treadwell:297 Nikolsky, G. V.:398 Nobiliey,, G.. Ky. :19'2 Norman, John R.:107, 411 Okada, Yaichiro:339 Okkelberg, Peter 0.:192 O'Malley, Henry:47 Ooster, John van:217 Osburn, Raymond C.:170 Ovchynnyk, Michael M.:225 Pack, Herbert J.:9 Parker, H. W.:303 Patten, William: 23 Perkins, C. B.:416 Peters, James A.:274 Peyer, Bernhard:200 Piast, tdi eR et: Pflueger, A1l:326 Pietschmann, Viktor:174 Plate, Ludwig:68 Pope, Clifford H.:343 Proctor, Joan:351 Raney, Charlotte F.:244 Ravenel, William de Chastignier: 33 Redeke, H. C.:120 Reed, Hugh D.:61 Reed, Roger J.:242 Reese, Albert Moore:216 Regan, Charles Tate:101, 410 Reid, Earl Desmond: 382 Reighard, Jacob: 340 Richardson, Sally Leonard:275 Rivas, Luis Rene:293 Rivero, Luis Howell:246 Romer, Alfred Sherwood: 283 Rosen, Donn Eric: 362 Roughilley.,) Ts C2193 Roule, Louis:102 Rust, Hans Theodor:146 Ruthven, Alexander Grant:226 Schachter, D.:228 Schaferna, Karel:151 Schindler, Otto:321 Schmidt, Frank J. W.:44 Schmidt, Johannes: 27 Schmidt, Karl Patterson:175, 280, 369 Schmidt, Peter J.:156 Schrenkeisen, Ray:50 Schroeder, William C.:284 Schulte, H. von W.:28 Schultz, Leonard Peter: 400 Scofield, Norman B.:182 Scortecci, Giuseppe: 409 Scott, Will:62 Seale, Alvin:314 Sette, Oscar Elton:232 Shannon, Frederick Albert:394 Shaw, Charles E.:229 Shetter, David S.:334 Shiraishi, Yoshikazu:335 Shufeldt, Robert Wilson: 38 Silvester, C. F.:10 Sancock, dx 4, sHawin’ H/..si62 Slevin, Joseph Richard:354 Smith, E. Victor: 89 Smith, George Milton:154 Smith, Hugh McCormick:260, 316, 381, 401 Smith, Mrs. Hugh M.:124 Smith, James Leonard Brierley: 330 Smith, Margaret Mary:312 Smith, Philip Wayne:267 Snedigar, Robert:205 Snyder, John Otterbein:103, 317 Soldatow, 1... Kemln Starks, Chloe Leslie:358 Starks, Edwin Chapin: 24 Starrett, William C.:230 Steinbach, José:13 Steinitz, Heinz:347 Stejneger, Leonhard:258, 259, 294, 295 Stensi6é, Erik:363 Stephens, Frank:63 Stevenson, John Alexander: 49 Steyn, William J.:221 Storer, Tracy 1.:417 Storey, Margaret Hamilton:190, 359 Stoye, Frederick Hans: 338 Strecker, John Kern: 420 Stuart, Laurence Cooper:247, 287 Sumner, Francis B.:318 Surbeck, Georg:130 Surber, Thaddeus:142 Svetovidov, A. N.:3 Tanaka, Shigeho: 336 T&ning, A. Vedel:176 Tanzer, Ernest C.:227 Taranetz, A. J.:113, 118, 12/5 Taylor, Edward Harrison:239, 286 Tchernavin, Vladimir V.:140 Tee-Van, John:250 Templeton, James R.:234 Terentjev, Paul V.:279 Thienemann, August:187 Thompson, William Francis: 328 Tinkle, Donald W.:346 Titcomb, John W.:17 Townsend, Charles Haskins:105 Trewavas, Ethelwynn: 243 Troemner, J. Louis:158 Turner, Clarence Lester : 333 Twitty, Victor Chandler: 344 Urich, F. W.:64 Vanderbilt, George:196 Villadolid, Deogracias V.:337 Vinciguerra, Decio:43 Viosca, Jxr., Percy:199 Vladykov, Vadim Dimitris: 372 Volsge, Helge:220 Wagler, Erich:162 Wakiya, Yojiro:85 Walford, Lionel A.:406 Walker, Charles F.:241, 408 Wall, Frank:144, 397 Walters, Vladimir:262 Ward, Jack A.:245 Watson, David:304 Weber, Max:54 Weed, Alfred Cleveland:165 Wehrle, Richard White:58 Welander, Arthur Donovan:370 Welch, Paul S.:186 Weller, Alfred:163 Weller, W. D.:14 Welter, Wilfred:90 Werner, Franz:78 White, Alfred McLaren:51l Whitley, Gilbert P.:367 Wiley, Grace Olive:138 Williamson, E. B.:29 Wilson, Henry van Peters:77, 86 Wolterstorff, Willy:65, 299 Woodall, Harold:104 Woodward, Arthur Smith:110 Wright, Albert Hazen:309 Wright, Anna A.:308 Wu Hsien-Wen:249, 422 Zaret, Thomas M.:248 Zhu Yuangding:371 ANNUAL LIST OF BIOGRAPHIES 1926- Jordan, Eric Knight (306). USO Cobb,asdonn N.w (5 380); Dybowsky, Benedikt (6); Forbes, 4 Stephen Alfred (7); Hay, Oliver Perry (11); Hine, James S. (12); Jordan, David Starr (292, 296, 387); Kishinouye, Kamakichi (8); Metzelaar, Jan (4); Pack, Herbert J. (9); Silvester, C. F. (10); Steinbach, José (13). 1931- Ehrenreich, Alfred (16); Jordan, David Starr (15); Proctor, Joan (351); Stejneger, Leonhard (258, 294); Weller, (14). 1932- Erwin, Richard P. (21); Evermann, Barton Warren (22, Johansen,-A. C. (18); Klugh, A. Brooker (19); M'Intosh, Ca C2i0)) Patten, Walilsiam! (23); Titcomb, sohn We s(57))) 1933—,ALeCock, iA. W..(30)2 Atkinson, N: J3:.°(25); Ayers, Howard 1934- 935 — TS3'6— 9 '3 "7 — 93:8 — WEE) 1940- 9 '4— Ola 1943- 10 (32); Johnstone, James (26); Maskell, F. G. (31); Ravenel, William de Chastignier (33); Schmidt, Johannes (27); Schulte, H. von W. (28); Starks, Edwin Chapin (24); Strecker, John Kern (420); Williamson, E. B. (29). Bacon, Daniel (35); Bensley, B. A. (34); Lataste, Fernand (36); Merriman, R. Owen (39); Nelson, Edward William (37); Shufeldt, Robert Wilson (38). Cunningham, J. T. (42); Hankinson, Thomas L. (45); Ishikawa, Chiyomatsu (40); Jenkins, Oliver Peebles (41); Knight, A. P. (46); Schmidt, Frank J. W. (44); Vinciguerra, Decio (43). Babcock, John Pease (48); O'Malley, Henry (47); Schrenkeisen, Ray (50); Stevenson, John Alexander (49); White, Alfred McLaren (51). Blanchard, Frank Nelson (376); Boulenger, George Albert (66); Clemens, Lucy Wright Smith (55); Giltay, Louis Pierre (59); Henderson, Junius (67); Hyde, Jesse Earl (52); Kampen, P. N. van (69); Longley, William Harding (53); McGregor, Richard Crittenden (56); Mosauer, Walter (57); Mukerji, Dev Dev (60); Plate, Ludwig (68); Reed, Hugh D. (61); Scott, Will ((62));\ Stephens, gerankm (6s) Uriech, ms Wie (64); Watson, David (304); Weber, Max (54); Wehrle, Richard White (58); Wolterstorff, Willy (65). Artedi, Peter (352); Berg, Leo S. (388); Borodin, Nicholas A. (70); Boulenger, George Albert (395); Lowe, John N. (dab is Embody, George Charles (74, 80); Goff, Carlos Clyde (73); Grinnell, Joseph (82, 88); Hoover, Barl BE. (72); Therning, Rodolpho von (421); Jacot, Arthur Paul (81); Kendall, William Converse (75, 79); March, Douglas D. H. (83); McMurrich, James Playfair (76, 87); Miranda Ribeiro, Alipio de® (8475 356)" Smith, Bo eVictor a(89)mwakiyal, Yior31ro) (8/5); Werner, Franz (78); Wilson, Henry van Peters (77, 86). Bass, Jn, ,00hnn F ((91));, Cope; bawand) Drinkers (C269), 3:57))); Lutz, Adolpho (93); Noble, G. K. (92); Welter, Wilfred (CIO) c Clark, Howard Walton (95, 96); Lahille, Fernando (94); Smith, Hugh McCormick (2607 316) 938i, 402) - Brumwell, Malcolm J. (97); Ditmars, Raymond L. (375); Reighard, Jacob (340). Allen, EE. J.9(98); Daniell, John Branklin, (99)));" Kelly, Howard A. (100); Regan, Charles Tate (101, 410); Roule, 1944- 1945- 1946- 1947- 1948- 1949- S50 its) e= 1953 — it Louis (102); Snyder, John Otterbein (103, 317); Stejneger, Leonhard (259, 295); Woodall, Harold (104). Bromund, Fred (108); Gage, Simon Henry (109); Juday, Chauncey (106); Norman, John R. (107, 411); Townsend, Charles Haskins (105); Woodward, Arthur Smith (110). Awerinzew, Sergius (114); Berezovsky, A. I. (115); Goodnoh, JE. Cli ftond Ay Giai6 )); "Morgan, Stanley Stewarte( lil); Reid, Earl Desmond (382); Sincock, Jr., Edwin H. (112); Soldatov, Vek. a (eli )ies Sumner, rrancas JB aaGs) 8) Tananetz, IX Mig ((abal3) 5 Desh Barbour, ) Thomas) (289); ,.Boulenger, (hag CG. h-(422);45Brimbey, Clement Samuel (281); Dahlgren, Ulric (123); Flower, Stanley Smyth (121, 396); Nellemose, (Captain) (119); Redeke, ‘H.-C. (2:20); Smith, Mrs. Hugh M., (124); Taranetz, IN (Cawley) Vie Allen, Weebe (lsu) 7 Allis, "dns, Edwazd® Phedps® (132) ; "Clark, Hubert Lyman (129); Cyren, Otto (128); Eigenmann, Rosa Smith (127); Hay, William Perry (126); Surbeck, Georg (T30)eS Woltexrstorfe, Willy (299). Bauman, Aden CC. (34/7) Camexvon, "A. -T. (135); Cockerell, T- DelA.nuits6) >) (Eager, (Grace (137) ;oEhrenbaum,., Emnst . (13/3); Wiley, Grace Olive (138). Hildebrand, Samuel Frederick (139); Tchernavin, Vladimir V. (140). Angel, Fernand (143); De Haas, C. O. J. (141); Hamilton, Rodgers: Diy (145). tHilidebrand, Samuel? B(3157 731974379), =3 837 384); Lonnberg, Einar (320); McIlhenny, Edward Avery (268); Rust, Hans Theodor (146); Surber, Thaddeus (142); Wall, Frank (144). Andersson, Lars Gabriel (149); Angel, Fernand (307); Berg, Leo Semenovich (147, 389); Bishop, Sherman C. (150); Blegvad, Harald (155); Eddy, Brayton (148); Ewing, H. BE. (152); Hoyt;,¢Southgate Y::.¢153)3 Schaferna,,Karel (151); Schmidt, Peter J. (156); Smith, George Milton (154); Wall, Frank (397). Barnhart, Percy S. (157); Bishop, Sherman C. (305); Engeling, Gus A. (159); Gowanloch, James Nelson (161); Libbert, Hans (160); Starks, Chloe Leslie (358); Troemner, J. Louis (158); Wagler, Erich (162). Babcock, Harold Lester (350); Ellis, Max M. (164); Muller, Horenz (313); Weller, Alfred (163) . 9) 5'4— LIS 5— 1956- IS) 5 7/= LSBs 1960- 19'61— LSI62— 19163 1964- 1965 — 1966 — 12 Duncker, Georg (166); Fassett, Harry C. (311); Gudger, E. W. (364); Johnson, Jr-, J. E. (267, 377); Lantz, “Louis A. (366); Weed, Alfred Cleveland (165). Allen, William Ray (169); Crimmins, Martin Lalor (168); Perkins, —C. B. (426): Bertin, Léon (172); Dunn, Emmett Reid (171); Gudger, E. W. (173); Hora, Sunder Lal (392); Osburn, Raymond C. (170). Dunn, Emmett Reid (378); Pietschmann, Viktor (174); Schmidt, Karl Patterson (175). Chabanaud, Paul (177); Gordon, Myron (178, 253); Maki, Moiehizro C183) Mayer, sh Rhez Cl] 9); Milvers Richard Di. (180); Nichols, John Treadwell (297); P£af£, J. R- (181); Schmidt, Karl Patterson (280, 369); Scofield, Norman B. (182); Seale, Alvin (314); Slevin, Joseph Richard (354); Taning, A. Vedel (176). Andrews, Roy Chapman (188); Hanko, Bela (184); Limbaugh, Conrad (322); Mann, William M. (189); Miller, Richard B. (185); Schindler, Otto’ (321); Storey, Margaret’ Hamilton (190); Thienemann, August (187); Welch, Paul S. (186). Crawtoxrd, = Sr, Donald) RR. Gl95))) (Cuntisy,. Braan. 1(94))); Mann, William M. (194); Okkelberg, Peter O. (192); Roughley, T. C. (193); Storey, Margaret Hamilton (359); Vanderbilt, George (196). Bruun, Anton Frederik (197, 323); Buen, Fernando de (324); Herre, Albert W. (198); Viosca, Jr., Percy (199). Alm, Gunnar (327); Anderson, Paul (201); Burt, Charles E. (405)') "Cope 7 Edward) = Drinkexee(29'8)));" Ege pe Vallinedm (325). Glauert, L. (202); Peyer, Bernhard (200); Pflueger, Al (S2B)re Breyer-de Rooy, P. J. (415); Chernov, Sergius Alexandrovich (20377 3418)" * Cook; ohannyerw Aj.) (206) "Grey, SMamaion (3)9/9!)7; Hartweg, Norman E. (204); Needham, Paul R. (207); Snedigar, Robert (205). d'"Ancona, ‘Umberto (208); Barnard, K: H. (209); (Clemens, Wilbert A. (210); Creaser, Charles W. (302); Davies, David H. (329); Davis, D. Dwight (211); Dymond, John Richardson (212, 385); Fox, Jr., Wade (282); Greenberg, Bernard S. (214); Howard, John K. (213); Miranda Ribeiro, Paulo de (215); Shannon, Frederick Albert (394); Thompson, William Francis (328); Wright, Anna A. (308). Baranov, Fedor Ilich (373); Fowler, Henry Weed (276); SIG = 1968- ISS) = USO IS) 7/d oe OZ — 3 = OTA — STS = S/O Is) 7 = US) ish 13 Ooster, John van (217); Reese, Albert Moore (216). Einarsson, Hermann (218); Michafowski, Jerzy (219); Twitty, Victor Chandler (344). Bigelow, Henry Bryant (285); Cochran, Doris Mable (301); Smith, James Leonard Brierley (330); Tee-Van, John (250); Volsge, Helge (220). Beaufort, Lieven Ferdinand de (264); Bragg, Arthur Norris (263); Cagle, Fred Ray (407); Hass, Robert L. (222); Klauber, Laurence Monroe (390); Matsubara, Kiyomatsu (331); Mohr, Erna (332); Parker, H. W. (303); Steyn, William J. (221); Turner, Clarence Lester (333). BUEZwGuadlhter —AGOlpho (223) Mitehell, EB. ud. .(224)): Ovchynnyk, Michael M. (225); Ruthven, Alexander Grant (226) eSchachter, Dian(228))) sShetcer, Davids. (334); Tanzer, Ernest C. (227); Terentjev, Paul V. (279); Wright, Albert Hazen (309). Frizzell, Don L. (231); Nakamura, Hiroshi (413); Sette, Oscar Elton (232); Shaw, Charles E. (229); Starrett, William C. (230); Steinitz, Heinz (347). Babbitt, Lewis H. (235); Eddy, Samuel (414); Landreth, Hobart (233); Peters, James A. (274); Shiraishi, Yoshikazu (335); Templeton, James R. (234). Hart, John L. (386); Herald, Earl Stannard (361); Hubbs, Carl lheavitec 10542) S365), 591) pe eLorcus His, aspera ds (l))i; Pope, Clifford H. (343); Romer, Alfred Sherwood (283); Scortecci, Giuseppe (409). Tanaka, Shigeho (336); Whitley, Gilbert P. (367). Donoso-Barros, Roberto (368); Ginsburg, Isaac (360); Gunther, Klaus (290); Meade, George P. (288); Mertens, Robert (300); Villadolid, Deogracias V. (337). Anderson, Jr., James Donald (419); Bennett, George W. (237); Conant, Isabelle (236); Cowles, Raymond Bridgman (266); Gaige, Frederick MacMahon (254); Gaige, Helen Thompson (254); Harrington, Jr., Robert W. (251); Koumans, Frederick P. (265); Lindberg, G. U. (2); Lutz, Bertha (345); Okada, Yaichiro (339); Schroeder, William C. (284); Stoye, Frederick Hans (338). Camp, Charles L. (417); Greenbank, John T. (240); Hansen, Donald. 16238) asNwkolsky,~eG. Ve. (39/8); Stoner, Tracy 1. (417); Taylor, Edward Harrison (239, 286). Se 1980- 9/8 A 9/82 LII83— 1984- 19/85 — 1986- S8i1= 1988- 14 Gloyd, Howard Kay (271); Hubbs, Carl L. (353); Martof, Bernard S. (255); Reed, Roger J. (242); Walford, Lionel A. (406); Walker, Charles F. (241, 408). Logier, Eugene Bernard Shelley (278); Maria, Niceforo (404); Tinkle, Donald W. (346); Trewavas, Ethelwynn (243). Raney, Charlotte F. (244). Eaton, Theodore H. (412); Haas, Georg (418); Johnson, Richard M. (349); Welander, Arthur Donovan (370). BoOhlke, James E. (374); Grodzifiski, Zygmunt (402); Rivero, Luis Howell (246); Stuart, Laurence Cooper (247, 287); Ward, Jack A. (245). Clay, William Marion (277); Cooper, Gerald Paul (256); Maslin, T. Paul (393); Zaret, Thomas M. (248). Blair, William Franklin (341); Kuehne, Robert Andrew (257); Lagler, Karl F. (355); Wu Hsien-Wen (249). Bredex, Urs, 1C.) M2 1G252)' /Goin, “Collemanmw (24/5): ) Myexsi, George Sprague (272); Richardson, Sally Leonard (275); Stensi6, Erik (363); Vladykov, Vadim Dimitrij (372); Wu Hsien-Wen (422). Carn, Jn., Archie F: (291); Czopek, Julmusz 9C403)'; Rivas, Luis Rene (293); Rosen, Donn Eric (362); Schultz, Leonard Peter (400); Smith, Philip Wayne (267); Svetovidov, A. N. (Die zhut Yuangdi ng: (Csi77))r. Hollister, Gloria (261); Miller, Frances Voorhees Hubbs (270); Smith, Margaret Mary (312); Walters, Vladimir (262). BIBLIOGRAPHY 1. Alexander, Richard D. 1974. Jasper J. Loftus-Hills. (3):812. N 2. Andriashev, A., and Z. Krasjukova. 1977. G. U. Lindberg, 1894-~ SG CS!) O12 6S), IN jaB,s 2 , E. A. Dorofeyeva, and N. V. Parin. 1987. A. N. Svetovidov, TISI0S-—T986.° (Gai27 2-2 713 Nie vse 4. Anon.) 1930-2 sJanyMetzelliaan, 618) 2al—119)2'9) 24 (2) 23-24 NB, 2 £EIBZOS JohnenN. Cobb. Gly 24 iN 30. shale USSOs TSB OF 1930. Ws) S}00)e ifs) 2i(0) - TSS OR aS IS'ON TSO" ALS) Siale AMS) SUE ULE) Bal ¢ GIS 2 UOISIZ) ISI2" USS r2" aS S2% ILS) S) 7x6 STS Zi IWS) 38h 6 IS) Bbc SSIS. IL) S3S}c POSS. AL's) 8} Sh6 IVS) S33} IL) 3\3)6 15 Benedikt Dybowsky. (1):24. Stephen Alfred Forbes. Kamakichi Kishinouye. Herbert J. Pack. Cc. F. Silvester. Oliver Perry Hay. James S. Hine. José Steinbach. W. "D." Weller. David Starr Jordan. Alfred Ehrenreich. John W. Titcomb. BN: A. Brooker Klugh. WoC. “Mi Tntosh: Richard P. Erwin Johansen. ((ale}) Grae (A) e622" (dEBE Barton Warren Evermann. William Patten. Edwin Chapin Starks. Nim Ul James Johnstone. Johannes Schmidt. H. von W. Schulte: E. B. Williamson. Aji We Alcock. F. G. Maskell. Atkinson. (3)) c25'6= Cove S6e (1):48. (2) iyi) (.4)pedi6:2.. (-Dy)ist 4:4). (2))/sat 2. (2) 112. (Ab) a18) 5 (CAS) BIS N (1) N (4)) 2161). N N (2) 2. (3):160. N 2 (4):166. N 4. (1):24. N, (3) eHE46 : N N N N B (3):160. N (ee) 2 74)8)- (A) 5s"418% (1):48. N N N ’ N B (1):47-48. SJAic 373). 347. SG 36% 375 Shehic 39). 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 5iOVe BIG Bee 53). 54. Bible 56). Bis UGE USS} Siar 1934. 1934. 1934. 1934. 1934. HOS 5k ISIS '5 7. 935% LS) SEY 985). 19 3'5 x0 19 /3)5)e S36. 1936. 19362 19316" 193'6< USS e 987 - USSF LOST UNS) Sh T/ 0 UOT 16 Howard Ayers. (4):228. N William de Chastignier Ravenel. (4):228. N B. A. Bensley. (1):60. N, B Daniel Bacon. (2):104. N Fernand Lataste. (2):104. N Edward William Nelson. (2):104. N, B Robert Wilson Shufeldt. (2):104. N R. Owen Merriman. (3):144. N Chiyomatsu Ishikawa. (1):48. N Oliver Peebles Jenkins. (1):48. N J. Tee CunninghamenG2)%-252 3 N Decio Vinciguerra. (2):112. N Prank: J.“ Wes Schmidityy1@3))i159 IN Thomas L. Hankinson. (4):202. N A. (PP. Knight).4 (4) 20 22°N Henry O'Malley. (2):132. N, B John Pease Babcock. (3):184. N John Alexander Stevenson. (3):184. N Ray Schrenkeisen. (4):244. N Alfred McLaren White. (4):244. N Jesse Earl Hyde. (1):80. N William Harding Longley. (1):80. N Max Weber. (1):80. N Lucy Wright Smith Clemens. (2):148. N Richard Crittenden McGregor. (2):148. N Walter Mosauer. (2):148. N 58. 59. 60 (Sle 62. bai 64. 657 66. thc 68. 69) Mie Halse 2 W32 74. Us Gy. Ue Uke Ue 80. 81. 82. 83. E987 LOST AS) 34/ 1937. WSS INS) Shthe MWS) BVT ISB) 6 UY 6 US)Si7/e iS) SNe UI SVS 19138" IS) ei}e LS) BIS)6 Ie ESESIOF. INS) SIS) ALS) 33) 4 ILS )3IS) US )3) 6 TS S.9% NS) SIS) 5 US) EIS) gS OF SISO 7 Richard White Wehrle. (2):148. N Louis Pierre Giltay. (3):200. N Dev Dev Mukerji. (3):200. N Hugh D. Reed. (3):200. N Will Seotes (3) 3200 ..N Frank Stephens. (3):200. N BE. We. Uraichaacs 3200 aan Willy WollitexzstorftE. (4)::.2/43i%7 BP George Albert Boulenger. (4):244. N Junius Henderson. (4):244. N Ludwig Plate. (4):244. N P. N. van Kampen. (4):244. N Nicholas A. Borodin. (1):56. N, B John N. Lowe. (4):210. N Earl E. Hoover 2 5(1'))3\63'..N,7 2B Carlos Clyde Goff. (1):63-64. N, B George Charles Embody. (1):64. N William Converse Kendall. (1):64. N James Playfair McMurrich. (1):64. N Henry Van Peters Wilson. (1):64. N Franz Werner. (2):122. N William Converse Kendall. (2):122-123. B George C. Embody. (2):123. B Arthur Paul Jacot. (2):123. N Joseph Grinnell. (2):124. N Douglas D. H. March. (2):124. N 84. 85. 86. 87. 88. 89. 0% Ales B25. 33). 914". 951. 96. SIC 38% We 100. al(@ake OF POS. 104. LOIS 106. LOT. 108. HOS". USBI) e IMs SISVic IS) Ss)c US) SS) c TS ESI9K. US} SI) 1940. 1940. 1940. 1940. 1941. 1941. 1941. 1942. 19 :4\3\- 1943. 1943. 1943. ISA SF. 1943. SAS \. 1944. 1944. 1944. 1944. 1944. 18 Alipio de Miranda Ribeiro. Yojiro Wakiya. (2):124. N, Henry Van Peters Wilson. (2 James Playfair McMurrich. ( Joseph Grinnell. (3):184. B E. Victor Smith. (4):244. N Wilfred Welter. (1):60. N John F. Bass, Jr. (2):148. Gs Ks Nobili 78. C4u2745275 .0'N Adolpho Lutz. (4):275-276. Fernando Lahille. G2) i224": Howard Walton Clark. (3):19 Howard Walton Clark. (4):27 Malcolm J. Brumwell. (1):62 Bi. J. “Alen > 4Glri68 Nn C2) a4 B ):124. 3) 31383" N 7 abe Ny Bee N Zi IN 8=27/9)e o }8 John Franklin Daniel. (1):68. N Howard A. Kelly. (GS) FCB N C. Tate Regan. (1):68. N Louis Roule. (1):68. N John Otterbein Snyder. (3) Harold Woodall. (3):204. N Charles Haskins Townsend. Chauncey Juday. ( 1):68. N John R. Norman. (3):196. N Fred Bromund. (4) Simon Henry Gage. 221616). IN (4):266. :204. N Gob BIS iS) ¢ N B B B, N N P 9 110. ___.. «1944. Arthur Smith Woodward. (4):266. N Dee LIAS Stanley siteware, Morgan. (1) :59'. N 2 eee 94 5. OE CWA nH oa mC OCK mune. G2, )rsele27 2 IN HOS eee O45 ).5 Al Rakanmecsr. (si) imleae).N 114. ___. «11945. Sergius Awerinzew. (4):239. N (but see 1947 (10) 8 B/S) 3) PS eee LIAS MA els VWBerezovsky =. (C4) 239)%. SN ieG eee l 9) 4 5 Clarif ond eA sGOOdnOhn, oir. C4). 239) 4 oN ely] LOA ony VK Oldatowes | (ANI 29h. oN Ino y eel 914 Oyen eA Olt lcd rlet zim (C4) e219. aN 119. __—_—«.~« 41946. Captain Nellemose. (1):54. N U0, 6 RSESo ils wo MEClOwGs ale S58) 21 eee LI4AG Stanley Smythe Blowers —@l))is56i7eN 122s LIAS. Bis Ge. BOUdenger= + (S))e:ilo2 . =N 12S L4G Ul rie Dahlarents —(@s)) shoss oN 124 LAG Mrs. sHugh eM (Smbtente (Sy) taste aN 25 4G Al STaranecz. (3) Le 4 aon 26 UIA Wallan Perry Hay. (ljt7/S). =v 27> ee LI4 75 Rosa Smith -Eigenmann. (1 )i75—76. «Ni <8 F216 ee SOAP OltatoreC yee = (ele) Neato aN 12 Oe 4p Hubert wm ymam Clarke (esh)ecee leat ESOS. L947 Georg (Ssurbeck. "(3267 N ES lees LOAN] Wie lute sAvloMemt.s | (F4))ese29)4beeNn 3 2 S40 Edward Phelps sAlelats Wits =(¢4?)t:) 29/5 oN 1S Sy ae. 194138 Ernst Ehrenbaum. e(l)75'2 N 134. - 1948. Aden C. Bauman. (1):76. N eS LIAS etn 1) Camenone clo)es 7 Grr IN P3i6.. SSITie 138) 139° 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150). Sse MSZ INSEE 154. SBE 1T5I6)- teWhs 158. L59)- 160. ilsabs 1948. 1948. 1948. 1949. TOAST £9510). 79/501. IES) 19/50. IS Wc 9S Or T9351. IES) ey U9 5K ise 1s )layabe Spite 9/515. IS) bs ALS aS 19S. SI5)25. USS) 7A AS) Se 952i Is A 20 T: 1D. A. (Cockernel Gl) e/6r. (N Grace Eager. (1):76. N Grace Olive Wiley. (3):231. N Samuel F. Hildebrand.(1):87. N Vladimir V. Tchernavin. (3):236. N G.. OF (GR) gia DemHaalsras (1s) 37d. aN Thaddeus Surber. (1):71. N Fernand Angel. (3):252. N Colonel Frank Wall. (3):252. N Rodgers D. Hamilton. (4):329. N Hans Theodor Rust. (4):329. N Leo sSi« Berg. sl) 112)., N Brayton Eddy. (1):112. N Lars Gabriel Andersson. (2):184. N Sherman C. Bishop. (2):184. N Karel Schaferna. (2):184. N H. EB). Bwings (3):j256.. .N Southgate Y. Hoyt. (3):256. N George Milton Smith. (3):256. N Harald Blegvad. (4):324. N Peter Ji. Schmidt. (4):324. N Percy S. Barnhart. (1):58. N J. Louis roemnerm. (1): 585 .N Gus Aw Engeling. {(15)):59)-, .N Hans Libbert. (2):125. N James Nelson Gowanloch. (3):221. N eal N62. 2). 1952). Brich Waglere. (4) 3289. N NW63ee cs. 1953. Alfred Weller. 002): 132, N 64... “1953. MaxM. Eldis.. (4) 254. N 165. __. 1954. Alfred Cleveland Weed, 1881-1953. (2):163-164. N, By 166%. -«-.. 1954). Georg Duncker!s (2!) 168 ON 6s wee 1954 Je Ee wiohnsion 7. or-eu(s)) 3.2467 N 68s sae 1955): Colonel = Maztin Lalior’ Crammins:.. (2:58 .. N 1692 =. : 1955 William Ray Alventyo(4) 3325. iN 170. = —: 1956: Raymond CC. Osburn scy34 oN L772: ==: 2956: Emmett” Reid Dunn.) (2 yeni 136 .. iN £72. jee: 29565 Léon Beatin.{ 102 ).ebsier N 73a meee: 1956. Bs We Gudger.) (2%). Tsien 742 aoe: 1957: Viktor? Pietschmann.7\(4) 2sSisN Spee LIS. Ke Pe Se himaditr.: 3(/4 Soran 176. _.. 1959). A. Vedel.Craning). .(dyxseo Ny. 77. a. 1959. Paul) Chabananudyc¢2)i:84..5 N 178. __. 1959. Myon. Gordoni. .(2)i:164-185 iN 1790 eee) L959. Era tizt Mayers, 4(72)) (28S oiN gO, 1. 1959. Richard PrOMiller..0 GZ) - 48 Sayin 1S 9592 J). Rice Poalter 8 (22) ) RBIS aN v2... 1959. Norman, i.i Scofiedaso( 2) L850 SS L959 osMomehuron Makal.9 s(Sire Zee. IN 184:> .:~« W960. Bella. Hankol.) (2/164. N 185. 7 960) Richard) Bf Madders 2\(2)): 64)-e IN 186. . 1960. Paul S. Welch. (2):164. N 22 187. ___. 1960. August Thienemann. (4):391. N 188. ___. 1960. Roy Chapman Andrews. (4):395. N 199. _ 7. 1960. William M. Mann. ((4):S95.. N L905 oe 1960" Maxganeits Hin Sironeyi@4)) i395. \N VO ee. TIGI Dr. Britain Cumiteasy mn @2piVZi6.0r0)N 192. 2). A961) Dr. Peter tO) 1Okkediberigims (2) )7::2/610)5 oN 193. _._.. 1961. Dry TSI. iRoughtey ane2261.5 N 194. ___. «1961. William M. Mann. (3):368 N 195). los. S61 Donalliahiky-: (Craw forxdiya Sia-5 ((4})/:i5a197F° ON 196. ___. 1961. George Vanderbilt. (4):519. N WS)o os BSISAAG Na ie feheroNbeyn ((Ib)) 8 Bisbee} OSS ee 9IG2 ) Al bert. Wea Herrera (el5)i2i5sl teen 1995 se W962.) Percy Valoscanwira-. (ls)kp2)515.) IN 200. Looe 19635 Bernhazd) (Peveraas)s223)7 N 201 oe LI63 Paul (Andexson-ea@2):4ri2n oN, B 202. —. 1963. Dr. LwGlavemte .C4)ii723'.0 iN ZOB. sa aSI6i4). Dre Si Ale Cherniovicnn())):2/55. IN 204. ___. 1964. Norman E. Hartweg. (1):256. N 205. 0. D964 Robert Snedilgarc: (())i256r Nig B 206. ___. 1964. Fannye A. Cook. (3):598. N 207. ___. 1964. Paul R. Needham. (3):598. N 208. — 9. 1965. Umbertor divAnconai: (cl) 124 -7)N 209% 2 UIGS\. Drs Key Woe Batanalaley aay) i 62/4 IN 210. 2 TIS. Wil bert AClemensi s (19) i:es2(4\-aeN 20 oe NIG SD) (Dwight DawaAls mel) i224; ENP eB DAU2y. . 1965. John Richardson Dymond. (1):124. N PS io 214. Za Si 216. 7 © 218. PANSY 220°. Dye. 222% PIIRESE 224. PUPS) 226. PIA T| « 228°. Re Zire PeSAL 232. SKE 234. 235%. PISS) 6 235i 238% IGS L965. IG 5. 1966. T9I6I6 . JEST e ISS cs 1968. T9/69.. LIIGS: 197/Ole ALS) 7/0) = ALS) 7/A0% IES PINS INS) 7b e LOW ly. POW 2). OR 2s USPS OR 0 US) Shc 19 3 UST Sc WSU USAT 6 19 7/80. Colonel John K. Bernard S. Paulo de Miranda Ribeiro. 23 Howard. Greenberg. Albert Moore Reese. (2):380 John Van Dr. Hermann Einarsson. Jerzy Michafowski. Ooster. Dr. Helge Volsege. Dr, Wisk ivam Je Dr. Robere i. Gualter Adolpho Lutz. Io Oh Michael M. Ovchynnyk. Alexander Grant Ruthven Tanzer Memorial Fund Madam D. Mitchell. ( Steyn. Hass. C2) Sel 3825 (2) 3503). (3) 3668 (242s 3) 598i N Schachter. (4):768 Charles E. Shaw. William C. oN IGS jr. PAZenuwle. Starrett. (i): 2016 aN (4):906. N Oscar Blscons Sete. (4) 906. Hobart Landreth. James R. Lewis H. Isabelle Templeton. Babbitt. Conant. (Gyis63 9% IN (CS) )y OS SiS, (21) etoile N N N (els Oly. N (1b) 8 ZOD HZ) George W. Bennett. (3):614. Donald F. Hansen. (2) S800 N N (3s) ES99%- N N C3) Sek (Cal) pebsyale N N C2) 2627 N N (AS BUSH N N N N, ‘ N N (43543. (2), 4.2254 2'3%. B ’ B N N (CARES ASals) 7a) B (2) S807 aN (Ernest €. Tanzer). (32588. 24 239% —. U8. Edward) Harrison) Tayjior-n(s)) 55). IN 240). _.. 19178... John, 1s (Greenbank. (4dEr 40! iN 2A. 2, UTS Charles sF 3 Walker (e262... N 242. 2, 1979, Roger, J.,Reed., (4)i 57. N 243. __. 1980. Ethelwynn Trewavas. (4):952. B 244. .. 1981. Chanlotte FF. Raney..(3)): 749. N 245 os, UBS) wack lA... Warde (lr 285i oN 2467 os U3 Luis Howeld Riverons (2): 582-583.) IN 247 oo. 983) Lawrence (Ce Stuaratre (3)! 853— 85425 N 2485 co 19/84. Thomas Mi. Zaretsn 4) 311042 5 N 249. - 1985. Wu Hsien-Wen. (4):1087. N 250. Atz, James W. 1968. John Tee-Van, July 6, 1897-November 5, UIG7 > es) \OCO—6617 aN Zoe. » 1977. RobewmeawW. Harringceon, Jr., LITT T975— \(s)e204— 20'5N, BP PABA 7 ELSI.) (Ce Ma eBreden- mois) 8977 — 19/833) 85S — i560 Niel, Pp 253r. and Donn E. Rosen. 1959. Myron Gordon, 1899-1959. ‘ C4) si 2-394.) (Bye 254. Bailey, Joseph R. 1977. Helen Thompson Gaige; Frederick MacMahon Gaige, 1890-1976. (3):609-611. N, B, P 255i . LOWS, Bernazd NS. Manto£. (2): 3 80—381'- Nj), B 256. Bailey, Reeve M. 1984. Gerald Paul Cooper. (4):1042-1043. N, B, P 257. Barbour, Roger W. 1985. Robert Andrew Kuehne, 1927-1984. C2) 521815: 2:Si IN eB aee 258. Barbour, Thomas. 1931. To Leonhard Stejneger. (3):73. P 25S . 1943. Leonhard Stejneger. (1):1. N, P 260. Bartsch, Paul. 1941. Dr. Hugh M. Smith, director of the Philippine Cruise of the "Albatross." (4):209-215. B 261. ZOZr 263% 264. ZO 5). 266. 267. 268. 269%. Zils UAL AY PAS 74 bs 274. 7 Se 21 Or Path ihe 2708) 25 Berra, Lim M. lose. (Gloria Holds ters 19100 -—19/8i8"= (4) ass. N, B, P Berry, bred. Lice. Vladimir Walters, O27 — 98776 14") 2 IIS 5 Wine tes TY Black, Jeffrey Howard, and Charles C. Carpenter. 1969. Arthur Norris Bragg, 18 December 1897-27 August 1968. (029) 4 LSE 74210 N By oD Boeseman, M. 1969. Lieven Ferdinand de Beaufort, 1879-1968. 22-2 NB 2 oie redertck 9p Kounans,, LIOS—N97 7. (cA) 802). Ni, By, P Brattstrom, Bayard H. 1977. Raymond Bridgman Cowles, 1896- WS) TS) com (CS) St SLAG ae 12h qe 12 Burr, Brooks M., and Lawrence M. Page. 1987. Philip Wayne Smithy a Loe — Leone (35) 659 —S4.0e0 NB, P Cagle, Fred R. 1950. Edward Avery McIlhenny. (3):245-246. N, B CaSey abe Cen LI 40m Cope] ine Mate) 2 OOo b> Chernoff, Barry. 1988. Frances Voorhees Hubbs Miller, Sahel (C2) be a Sohne Whi ealsbio 12 Clay, William M. 1979. Howard Kay Gloyd (1902-1978). (iby) SALES) g yg wtp ele Cohen, Daniel M., and Stanley H. Weitzman. 1986. George Sprague Myers, 1905-1985. (3):851—-853."N, BB, P Colbert, Edwin Hee lI8G. "Coleman. Goin). 1 9Hel— 19 8ior (4 eo at 04 3e Ny By =P Collette, Bruce B. 1973. James A. Peters. (2):388-390. N, B, Pp 1986. Sally Leonard Richardson, 1944-1986. (4):1043- 10/4'50 NB 2 Conant, Roger. 1966. Henry Weed Fowler, 1878-1965. (3):628- D4Sloe ANpe i, 12 MICA Wineleramne Mar TON Clic, a 06 — 196s C2) O16) 3s Ni, Crossman, E. J. 1980. Eugene Bernard Shelley Logier. (S)))5 UPA ClG. Wipe yp 12 Zi. 280. 281. 282. 283%. 284. 2Z85\- 286. 2801. 288. 2897. 2907 Payal 2912). 293i DIA 295i. 296i. PESTLE 26 Darevsky, Ilya S., and Leo I. Khosatzky. 1971. Paul V. Tezentvev, (1190S = 1970) C2 S182 S64 eeNE BP Davis, D. Dwight. 1959. Karl Patterson Schmidt, 1890-1957. (BYR UG GLI aisle 1D Davis, Harry T. 1946. Clement Samuel Brimley. (3):182. N, B Dessauer, Hic.) 1965) Wade hoxpardias me! 9:20 —A9 GAR eileen 2 310 NI B, P Dick, Myvanwy M. 1974. Alfred Sherwood Romer, 1884-1973. CI ESS 2.94 aa Nia 8 UOT Wiliam Ca Sehroeder.. 189/419 7 2g(2 is 42 4 esis Nb. , and William C. Schroeder. 1968. Henry Bryant Bigelow, 87 SAS 7) aee3)) 2 Oy) OO ON eS ee Duellman, William E. 1978. Edward Harrison Taylor, 1889- OT Brn) el Sila 3 CukIN) aS ee, 1983" Laurence, Cooper Stuart. (4):11t6—-1118. B, P Dundee, Harold A. 1976. George P. Meade. (1):220-221. N, B Dunn, Emmett R. 1946. Thomas Barbour, 1884-1946. (1):1-3. N, By, ee Ebeling, A. W.: 1976. KiaussGunther, 1907-1975. 514) 2845. N78 Bhrenteddy. Davide. 3 8i/pwARenTe Lh .. Carr mdr. pa LOSS LSS iii (4) 31087-1089 . Nae Beale Evermann, Barton Warren. 1930. David Starr Jordan, the man. (40 SS =0)5)0 By ee Fink, William L. 1987. Luis Rene Rivas, 1916-1986. (1S) i222 6:92 slew Nip Bae Fisher, Albert Kendrick. 1931. Leonhard Stejneger. (3):74 B35 8), 2, 1943. Leonhard Stejneger. (3):137-141. B Fowler, Henry W. 1930. David Starr Jordan and ichthyology in the, Paci fic. » (4) 2108-109)" )B 1959. John Treadwell Nichols, 1883-1958. (1):83. N, B Z2918% 299%. 300. SHOEF 302. 303. 304. 305i. 306. 307 308. 309. SNe Shihe S22 Spisip 314. SHS sally SiG 27 19163. Cope in’ retrospect. (1) 2195-198. B Breytag, Gunther. .1947> Dr. Willy Wolterstortt. (3)):215. N, B Gans, Carl. 1976. Robert Mertens, 1894-1975. (2):420. N, B Goin, Coleman J. 1968. Doris Mable Cochran, May 18, 1898- MES? P72 GIGIAGIE (Si) SSGNSSSA. se tsi 12 Gorbman, Aubrey. 1965. Charles W. Creaser. (3):398-399. N, B Grandison, Alice G. C. 1969. H. W. Parker. (2):416-417. N, By, .P Gregory, Wa Ka S9s7e DavideWatson.o(3) 2197. BPePe Grobman, Arnold B. 1952. Sherman C. Bishop, 1887-1951. (SN BUA YH=UAS 1, 12 Guerard, Albert. 1926. Eric Knight Jordan, 1903- LIZ6 RL S2) suppl. NP 8 Guibeyi hI 5is RFemnandsAngel,q18é61—1950 eo (1)st=2. B, P Hamislton,s JGe ,aiW.. Win 29165). aMrs arAnna BA. eWwoight. (1) 31 245eN, B 1971. Albert Hazen Wright, August 15, 1879-July 4, TSAO OPA) BSH PSSA INTE ® sl | fe! Hanna, G Dallas. 1932. Barton Warren Evermann, 1853-1932. (GAD) ce iGel— 62) BP 1954), Hanry Coy Fassett. (2)2167% N Heemstra, Phillip C., and Michael N. Bruton. 1988. Margaret Wesesr Ghimelp WS SSS 7/5 CY SA Sinaia MWip V2}5 0 12 Hellmich, Walter. 1953. Lorenz Miller, 1868-1953. (3):133- 13.4. IN, sBiak Herre, Albert W. 1959. Alvin Seale, 1871-1958. (1):85-86. N, B Higgins, Elmer. 1950. Samuel F. Hildebrand as a government Semeniteasity ae) si8i—ael 3B Hildebrand, Samuel F. 1941. Hugh McCormick Smith and the Bureau of Fisheries. (4):216-220. B, P Hubbs;,, Card L). 1943). John O. Snyder® (4): 265-2665 BRB, P Sus) c S\7A(0).6 3:24. S226 323%. 324. 325). 326. S21. 3)2/8r. 32955 330. 331. 3I3:2%. 333% 334. 335) 336%. 3378 S''Gi- 33 9r. 28 1945. Francis B. Sumner, 1874-1945. (4):183 ___- 1950. Samuel F. Hildebrand, ichthyologist an hezpetologisit: * (1) 112-045 SB,; 2 P 1950. Einar Lénnberg. (1):70. N E60 Dr. Otto Schindilexrts ((Ss)i:269. Ni, B 1960) Conrad Limbaugh. (S)ij269—2710).. Ni, Bi, Pe 1962. Anton Frederick Bruun. (2):481-482. B,; 1962. Fernando de Buen. (4):875-876. N, B, P L963. Vilhelm. Egesm ()i2220-222). ON) By SP), 1963. Ale Pflueger. (@y)s223r N 963. Gunnar Adm.” (1) 32245 N 1965. William Francis Thompson. (4):542. N, 1965.5 1IDavid shia sDavéiels .. c(t): 543... IN, B - 1968. James Leonard Brierley Smith, 1897-196 G60. NEB, Pe 1969. Kiyomatsu Matsubara, 1907-1968. (2):42 Bye 1969. Erna Mohr, July 11, 1894-September 10, (3) 716416." Ne Bi OP 1969. Clarence Lester Turner, 1890-1969. (3) N, B USL. SDaviades seShetteryuel OL O—19i7 Oe Gia) ys) STAN 1973. Yoshikazu yShurweanshin. (C2) 393 —s9 45 Nip es 1975. Shigeho ‘Tanakay,, 878-1974. 1(4)) si7/S)27eaNi, 1976. Deogracias V. Villadolid, 1896=1976- ( UST7? Ss Prederick Hans *Stoyey,” 18s8v—ao7dise (3h): B, P and Katsuzo Kuronuma. 1977. Yaichiro Okada, 1206-2074 0N, BpoP Ms —s (1 =84. N,. 8, d P B 8. (3): 659- 0-422. Ny 1968. 2646-647. N B, P Sr 6! 7 aN 615-616. N, ILIA S IS) IS e 340. 341. 342. 343. 344. 345. 346. 347. 348. 349. S500. 35s. SDZh 55S 354. 355). B5'6r SS 29 , and Karl F. Lagler. 1942. Jacob Reighard, 1861-1942. (ip) 63=64 "Nes ee Hubbs, Clark. 1985. William Franklin Blair, 1912-1984. (2):529-531. N, B, P Hubbs, Laura C. 1974. List of students whose PhD degree was under the chairmanship of Carl L. Hubbs through 1974. (3) 609 —c00r Inger ,/eRobereer.. 9745 ClrefEoraieH. ‘Pope. 1/4 )isLON2YEN, B Kelly, Douglas E. 1967. Victor Chandler Twitty, November 5, 1901-March 22, 1967. (4):887-888. N, B, P Kennedy, J. P. 1977. Bertha Lutz, 1874-1976. (1):208-209. N, B Kluge, Arnold. 1980. Donald W. Tinkle (1930-1980). (3):572. Ni. BP Kwehnne,; Robert, A. 19725 Drs Heinz Stentz, 1909-971. (3) 2 G0S—G6il0 =Ni,- *B), =P Leviton, Alan E. 1964. Sergius Alexandrovich Chernov, 1903- E964. (2):3466.°B,"P Liner, Ernest A. and Allan H. Chaney. 1982. Richard M. Johnson. (4):987. N, B Loveridge, A. 1953. Harold Lester Babcock, 1886-1953. (3)) 41 34=153'5\> 4N/7 9B Mann, William M. 1931. Joan Proctor. (4):166. N, B Merriman, Daniel. 1938. Peter Artedi-Systematist and Vfehthyologisit.9 (A)ES3S=3989 B Miller, Frances H., Clark Hubbs, and Earl L. Hubbs. 1979. Carl -GP*Hubbs’. (4) 367/56). @N 78 Miller R. C. 1959. Joseph Richard Slevin, 1881-1957. (1):84- 85. N, B, P Miller, Robert Rush. 1985. Karl F. Lagler, 1912-1985. (4))) SILOS @—Salojsale MiG zi, 12 Myers, George S. 1939. Alipio de Miranda Ribeiro. (3):184. B 1940. Cope as an ichthyologist. (2):76-78. B, P 369. 37 Or S)f/ lic 372). S13 37/4... 3/5. B/G. 30 1952. Chloe Leslie Starks. (2):124-125. N - 1961. Margaret Hamilton Storey (1900-1960). (2):261- IASG Np 1p Ie 1976. Isaac Ginsburg. (1))/3217-219. N,°B, P , W. I. Follett, and William A. Gosline. 1974. Earl Stannaxd yer aiahji( 1 )¥: 29:12:92) aN eB co Nelson, Gareth, James W. Atz, Klaus D. Kallman, and C. Lavett Smith. 1987. Donn Eric Rosen, 1929-1986. (2):541- 546 Ni, By Py , and Mee-Man Chang. 1986. Erik Stensi6, 1891-1984. (2) 25.5'8i5 Nie IB) ae Nichols,) John! T3954") EB. Wee iGudger.. (2)M64—165. B Norris, Kenneth S. 1974. To Carl Leavitt Hubbs, a modern pioneer naturalist on the occasion of his eightieth year. (3)i: 53-5 9AR Bye P Parker,® Hi. Wi) 19154 2 Louis sAresbamtz’. 4 (179 aN Paxton, John R., and Douglass F. Hoese. 1975. Gilbert P. Whatley, US903-—19 75) (4) ia 9i2e95). ON, BB), ae Pefaur, Jaime E. 1976. Roberto Donoso-Barros. (1):219-220. Nie Bee Peters, James A. 1959. A bibliography and index of Karl P. Schmidt's papers on coral snakes. (3):192-196. L Pietsch, Theodore W. 1982. Arthur Donovan Welander (1908- WOSZANG (SEUSS o Whe 13 Qingwen, Meng. 1987. Zhu Yuangding (Yuanting T. Chu), 1896- S86) (CS )iI8S8—839 Ni Smee Renaud, Claude B. 1986. Vadim Dimitrij Vladykov. (2):559- 560he Np eB Ricker, W. E. 1966. Fedor Ilich Baranov. (1):147. N Robins, C. Richard. 1983. James E. Bohlke. (3):850-851. N, Bye: Ruthven, Alexander G. 1942. Raymond L. Ditmars. (2):131. N, B Schmidt, Karl P. 1937. Frank Nelson Blanchard, 1888-1937. 31 Gar 439/—15.00 Ni Bye Sialire >; 1954. J. Ep. sgohnson, Jnr.—- An appreciation. (3)3247-—248. 37.805 . 1957. Emmett Reid Dunn, 1894-1956. (2):74-77. B, P 379. Schmitt, Waldo L. 1950. Samuel Frederick Hildebrand. (1):1. P 380. Schultz, Leonard P. 1930. John N. Cobb, Dean of the College of Fisheries. (2):50-51. B 361. - 2. 2941. Hugh McCormick Smith. (4)2194-—209). N; B, By L 382. ___. «1945. Earl Desmond Reid Retires. (2):128. B 383. 2. L505 Samuel Frederick Hildebrand. (21)32—-7. By, P 384. , and Ann Shepard Green. 1950. Bibliography of Samuel F. Hildebrand « 1(1))k: 15198... ie 385. Scott, W. B. 1965. John Richardson Dymond, 1887-1965. or SSI6—ssist Bea e SCO elo A Onn Hart, L904 — 1917 si (2) 579 N), 8 387. Seale, Alvin. 1930. To David Starr Jordan. (4):106-107. B, P 388. Shapovalov, Leo. 1938. Leo S. Berg. (1):55. B, P B89e se. 1951 shea Semenovich Beng jaw! 876-1950 9a( 3) )ilesi5—18'6.. Biase 390. Shaw, Charles E. 1969. Laurence Monroe Klauber, 1883-1968. (Ae ceySeialsy 5 Ile eye he 391. Shor, Elizabeth N. 1974. Selected bibliography of Carl Leavitt hubbs rom 9m5 tor 1974 (3): 594—609)-, 392° Silas, E. CG. 956.5 (Sunder Wall Hora. (2) 384-1360 .N, B, P 393. Smith, Hobart M. 1984. T. Paul Maslin, 1909-1984. (3):806- S07) Np Be 394. , and Findlay E. Russell. 1965. Frederick Albert Shannon, SZ \5re) (4) 54a — 5427 ON, By, oP 395. Smith, Malcolm. 1938. George Albert Boulenger, 1858-1937. (15) 5). — 35.5 By, =P 396. « L946. Stanley Smyth Flower, 1871-1946. (4) 2185-187. B, ie) Silo 39/8). SIS)S) ic 400. 401. 402. 403. 404. 405. 406. 407. 408. 409. 410. 411. 42" 413. 414. 405%. 32 LOS Prank Waly) VS8i6s— 1S 50m (2)) ils — rR Soin, S-- .Ga,.and M. Vo Mina 2978. (CG. Vi. Nikolisky (Glgh0— STU Mio CAMB BUN a INi5 ll, 1 Sonoda, Pearl M., and Robert F. Inger. 1964. Marion Grey. GS) S9'8e" "Nie B Springer, Victor G. 1987. Leonard Peter Schultz, 1901-1986. (Els) 27 22 NB ee Stejneger, Leonhard. 1941. To Hugh McCormick Smith. (4):193. P Szarski, Henryk. 1983. Zygmunt Grodzinski. (3):851-852. N, B, P 1987. Julvusiz Czopek, 1922-19386. (3) 783Sr IN Tamsitt, J. R. 1980. Brother Niceforo Maria. (4):952-953. N, B Tanner, Wilmer W. 1963. Charles E. Burt. (4):722-723. N, B, P Thompson,, Paul» Ey.” 19'79'.9 Livonedy vAy.. Wal ford. (3) 75645 N Tanke. Donald W. 1969. “Pred Ray Cagle, 1915-1968.) (is) 2:20 N, B, P 1979. Charles F. Walker. (3):563+564. B, P Tortonese, Enrico. 1974. Prof. Giuseppe Scortecci, 1898- OH Sha (Al) StZ94SN YB Trewavas, Ethelwynn. 1943. Charles Tate Regan. (3):202-204. B, P 1944. John R. Norman. (4):265-266. B, P Trueb, Linda. 1982. Theodore H. Eaton. (1):245. N, B Ueyanagi, Shoji. 1972. Dr. Hiroshi Nakamura, 1906-1971. (GIS) S 321042 0lSi.= (Nyaa b Undemhill, James. 1973. Samuel) Eddy. (2)):392-393- N,. By, 2 Van der Feen-van Benthem Jutting, W. S. S. 1964. Mrs. Dr. P. J Breyer-—de Rooy. (3):598. N, B 416. 417. 418. 419. 420. 421. 422. 33 Wade, Wallace B. 1955: CC. 8B. Perkins, 1889-1955. (4)3323- 324. N, B Wake, David B. 1978. Tracy I. Storer and Charles L. Camp. Gil BIS S=IUs) 7/5 13! Werner, Yehudah L. 1982. Georg Haas, 1905-1981. (2):491-493. 1\ ee = ee 2 Wilhoft, Daniel C., and Keith A. Hawthorne. 1977. James Donald Anderson, Jr. (2):412. N, B Williams, Walter J. 1933. John Kern Strecker. (1):48. N, B Wright, Stillman. 1939. Rodolpho von Ihering. (4):244. B Viv Cnien Loco. WHS wen wen). 900 — 1918 5<) 102): 5oui—5o8 ene a rf a i ay ; re 5 % : o , 4 7 ani low ae cesesiay eer ebar ta @) «5 2G80L .8 odei law cls ah Prank weidy Speetstis- 1148 a gever vanbt (aden ora ie ral nent wD fotnsdn, t200L \toarebaA Bienod amet ores Ca¥ gtmBt LAW: 20 i | 404, see ; 7, ml ai. » paged n08-odl East tee \2dplaM: uth e Bex t saab _— a SRE ptea (SP ‘Teeet-o0er avenpAen vv .25°l .ngdd aly a Ra: 3. “we. 3 Sal iusz Ceopely 19h 1}: 0 oe 90, ra ‘. 1900; brothel _ pepeese-osay: 4 j a 4s, tp vee er om me yee 19a My . 74 + ra Oe | } ues | 79, 4 ee 3):564.°8 85. Meson 7 i . 407. i. Doni #, 1969. FES RAY 014, Un@erhit\, dames. 1872. ave} eh, 0° ou 2-909 P. 415. Van Gee Peeneval airy hia ‘3 ae Or. anh de t, Breyer-d# Rowy (21:38 We we ane oe Nie a A KEY AND CHECKLIST TO THE NEOTROPICAL SNAKE GENUS LIOPHIS WITH COUNTRY LISTS AND MAPS JAMES R. DIXON Department of Wildlife and Fisheries Sci Texas A&M University SMITHSON HERPETOLOGICAL INFORMATION RVICE NO, 79 1989 SMITHSONIAN HERPETOLOGICAL INFORMATION SERVICE The SHIS series publishes and distributes translations, bibliographies, indices, and similar items judged useful to individuals interested in the biology of amphibians and_ reptiles, but unlikely to be published in the normal technical journals. Single copies are distributed free to interested individuals. Libraries, herpetological associations, and research laboratories are invited to exchange their publications with us. We wish to encourage individuals to share their bibliographies, translations, etc. with other herpetologists through the SHIS series. If you have such items please contact George Zug for instructions. Contributors receive 50 free copies. Please address all requests for copies and inquiries to George 2ug, Division of Amphibians and Reptiles, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Please include a self-addressed mailing label with requests. INTRODUCTION The genus Liophis currently contains 35 species and 62 recognized subspecies. About 137 names have been proposed, attesting to extensive qualitative and quantitative variation between and among the species of Liophis. The purpose of the checklist, keys, and country lists is to offer museum curators and herpetologists a means to identify the species and subspecies of one of the most commonly encountered xenodontine snake genera of the Neotropics. Species lists are constructed from specimens personally examined and verified by me. The keys resulted from an analysis of variation of 5,198 specimens of all recognized populations. Some species have been recognized since 1758 and others as recently as 1987. Some species are represented in collections by only a few specimens (L. atraventer, L. problematicus), by several hundred (most species), or over a thousand (L. miliaris, L. poecilogyrus). Taxonomic data are generally adequate for most taxa. The checklist contains the accepted name and its author, followed by primary synonomies, their author(s), date, page number, and type locality. The primary synonomies are followed by the author, date, and page number of the first proper usage of the epithet, if necessary for clarification. Species are arranged alphabetically. Subspecies are also arranged alphabetically under each species, except for the nominate race, which is placed first. Synonomies for subspecies follow the accepted name, arranged in alphabetical order, followed by the author(s) name(s). A statement of distribution follows the synonomy of each species and subspecies, along with a citation denoting a publication with a distribution map. anol IaTuqog Bastagoses (Te ‘Yo Rnamioeqz 827.2 Fo ha need sved sacar eee say SST Yam 107 sisupsbe onthe i 2nots2al ) Bia ra 2 iT ea ft v -. ret St iter’ Anamptads 89. (2) inatruceiame. ce imutere receive $0 free goctees, Pleéoe athir@ee 411 fequente far regies and ingvistes to George Digs Diviaton of Aaphibtane and (led, Hationsl Mubevue of Mature) @ Wietery, Gamithecnien Institut{on, Woshinarany 8.6. 20660, U.5.A Please tuicluce » self-sddreanpd was) tha label with reqduaeta. 10. Uke 12. 13% KEY TO SPECIES OF LIOPHIS Posterior dorsal scale rows at least two less than at MiabOdY %2)..25 eheea hee Pete eEGee Eps «etrenGs the olesmebese Dorsal scall GynowSmwithOur seGUG EON, o.c15s 5-07) c1.070 010-010-0001 or cho one etait DOSall MS CallennOw sy ell 55 acl wonseeycrersrce cee torah av ev oi es ov h.e 6: cicaret ancrevarsrors GDR MRI. Doysaligscaletrows, 17-17-17. ......Stees cnet binow :8%6 800.2 SRESS three! supratabial’s ‘entering Orbit Iy.352 Jon. Sas OM. andinus Two supralabials entering orbit. reqinae (=oligolepis of others) Eight supralabials (rarely seven); dorsum olive green with or without reddish vertebral stripe and smal] dorsolateral DllacksSpOts ateas «fh biw o3de6 5 weedeat osanat jaegeri Seven supralabials; dorsum tan or brown with darker blotches and four blackish posterior lines, and a black iedged ;) whiter labial) stripe. scsccs. f tex ciate ges williamsi Ninetecnemidbody 'dorsal/’scall@ VOWS OF. 5 ..6.s ec cee cscs pesccwcsce Seventeen midbody dorsal SCal@svows PiO2. occ. cc wccsoedacwordneds BGPSdIMSCANER OWS 20 maxillary teeth .......... longiventris Venter with checkered pattern of black and red or yellow ....... Venter never checkered with black and red or yellow, but occasionally with black marks on lateral edges of Venbral's (sor ehocgr ores suias soa ee See! Sheesh. PR Rds lowtatscs Black lateral posterior stripe present and extends GncGeratl J sU2h 22 O¥a, WSO. SOT eS. Logs Rae a alto. Bae. tweds. NOGadS:GOOVE Ms <50e IRS. Sat eels Res Als Ole Belae-6.c go we wile acters Apical uscalewpitypresenth. os. vsoDett. dass: 598 Fe O40 8AGogy. wovoe: Nodscalle prtrrasaga ra: .ng84ns v9. ns9qn- awd fa-eomanah~ 2 FS taeniurus Posterior lateral black stripe occurs as an edge between 3rd and 4th scale’ rows G.....sceceeeesccsctoes reqinae Posterior lateral black stripe occurs over most of 3rd, one-third to all of 4th and occasionally on 5th scale NRW t feta a sotlen ey wiratar at nyo cseterrclei eis Ser six)-ova oie sual eva’ wie HtaIw # ee epinephelus MC Cail AV era eptctaee © rey clapevs cious: araieiae elbinveve fod olaiorels iste Sie Gen aaeseras TREO) SPST FENCE festae mo Pw 10 19 11 13 12 14 14. 1S. 16. Ly: 18. 19. 20. 2 22: 23). 24. om Ventralis<179) occ c5 5 HAPS ERAS TR RRO FOR Ries os cs scecictoneneteerseins Ventralls S186 |. 5ccos.d selene scsi sis obi eiele ala isats ateyetoters de sareelare frenatus Apical scale pit present, tail length >26% of total MONGEH: win e505 4 05 ws AT eS ORR oie sole Rites ctotehs © Scale pit absent, tail length <23% of total length ......2.....: Subcaudal's: >743+ >26: maxi] lary. teeth... .\4525%%7. SNR 2A juliae Subeaudalis <723 <2? maxillary teeth -....2050..5--» . epinephelus Banded dorsal pattern, occasionally a combination bands and reticulations; ventrals and subcaudals always checkered with black and red or yellow ...............eeeeeee Salt and pepper dorsal pattern, obscure in Amazon populations; venter light with some dark edging in all populations except Amazon, where large black checkered marks are present; Amazon population usually with dusky throats and white subcaudals, subcaudals of other populations white or dark edged ......... SEES RASERDRERETSES SSE E EE ESS cee eee Ae ee Ren miliaris Light body bands widened laterally into broad triangles ........ Se a ee ee ee longiventris Light&bedyLbands- of-equal-width: ..2<025 055. 55255..2087 5 cobellus Lateral -edge: of: ventrals«unmarked:s..<2.2..025%. 545.08 2790% 08% Lateral edge of ventrals marked with black ...................-. Dorsum with or without posterior lateral black stripes; S13 LMacd lary: teeth: «2% 6255.8s0 es ese eae se 2 eT Re. Dorsum with three dark lines from head to tail; 21 maxillary teeth; tail length about -26%: of -total-length:74; dorsum uniform color or variable, but never -green or olive green «ssscsscsccs. 3 SOSFI8. 259. 2805. 188: Subcaudals <73; dorsum olive green or green, with or without reddish dorsal stripe and small dorsolateral black spots ... 2029. 98. 28. STeI20. Folge 5. PORIE. SS ISS e: jaegeri Ventraiis: <1l6775.. 39. 2E08. Jeo. Se as Tae Aaa AS SR aes eS juliae Ventrals >782°92. 925. 08. 25 L200) S8208. ORS. Os 8. Ie. SSR. perfuscus 15 16 7 18 20 25 21 22 23 24 reel 26. 27. 28. rane |e 30. 31. one 33. 34 35. Dorsum without darké Stripes: «': e can 04). ARTS SER. US RS Dorsum, With dark SUrmiPese siete. ie es teleliere's weal ae OR TL ARIE Lai lealengtht>26%. of total lengthy <<<. ccscecds che SRS Ue RIG Vail length’ <23% ofi total length 12% cea ee Ee TS miliaris Subcaudal's *<93 Wits 0s AT EERS eae fom sate ete oe eae es Ue ornatus SubGaudalis S9Grhse* . RECS ERS CoRR SS ERRMRQGNSE SoG S SEW TIQ cursor Dorsum with three dark and two yellow stripes, yellow SURIPESTDEGIN ON SNOUL <..cccceccese csc ees so sss flavifrenatus Dorsum with three to five dark stripes, yellow stripes GOSONG rete c sac REAR Sete oe oS TOR FeO S Pee eee et bh triscalis Dorsal scale rowsvl9-W9=170% 2 28 Sos, FE 084. RA TR ARE SU TTY Dorsal ‘scale’ rowse19=19-15°0r7 19=19-93S 2780.2 .99 604.2 ORS. Dorsum uniform green or olive green, occasionally with an ill-defined reddish brown mid-dorsal stripe .............. Dorsum variable, but never uniform or with an i1?-defined/redditsh'mid-dorsal*stripe 9203.25.52: 2206. 288278) Ventiaerense "So. 258. cba Meee leo heme di eatistsacae ss cheat chabecds VENERANS eS LOO tee coe shee e een. PR ae el ts 4d” be eee viridis Venter rose or red; >62 subcaudals; >25 maxillary teeth BCI Ree SRE bd: Bene ey. maryellenae Venter black; <56 subcaudals; <24 maxillary teeth So tine a6 ees oc wel tote d She She cereals cit tA ave tls Detets S eee es atraventer One“preocular™. BUSTERS. Lae, Ale RAPES Sa oes AS Ol 79l S98. OL IW. 2788, TWO (PrEOCUars PNFLFIN 228. Gs FIFI HALTS SNS: 22S8250 IRI 82. festae Dorsum uniform brown or with two to five dark lines on aytan to /vight ‘brown ground -colior see 0F4289 05). 2899. 81822. 1! Dorsum gray or olive, with or without dark irregular marks scattered throughout; or dorsum brown to dark brown with large blotches or bands anteriorly ............... Dorsum tan or light brown with three to five distinct dark stripes, the median stripe beginning on the SMOULMOV ENes MAPOache estes ate so-c G Reretate accel aie Ga dvs wales Disiwiev Se acaneves a aoe Dorsum uniform brown, with or without dark freckles and a IdaMVieritrolateral. Wine «.-..cceereereretoreteeteteteretelere’e sagittifer 30 39 31 33 32 34 35 37 36 36. EV/6 38. 39. 40. 41. 42. a Dorsum tan or light brown with three distinct black lines, the median line beginning on the snout, >159 ventrals; >77 subcaudals; tail/total length ratio (males) >.24 .......csessseeecceeecevececscccs lineatus Dorsum light brown with a broad dark median stripe, narrow dark dorsolateral stripes, and dark ventrolateral stripes; median stripe begins on the nape; ventrals 133, subcaudals 36, tail/total length ratio .185 (known only from the male holotype) Sco no alg oe cM PE « sR 6 Ss & WOE ERY Che fete ae ee problematicus Dorsum gray or tan with darker blotches, no black dorsolateral line posteriorly; faint to moderately distinct dorsolateral light line posteriorly; dorsal surface of head gray, tan or brown with or without a whitish U,V,X, or Y mark on the parietals with exterior black edging, the mark may extend anteriorly to the internasals; with or without one apical Scale Pit ....5.ccscseensmecessecssccecscewerccceesees Dorsum dark brown or chocolate brown, banded or blotched anteriorly with contrasting dorsolateral black and white lines posteriorly; no apical scale | Gas Hee A RARsIAIEN Sian eclain Heke ei onic cS Adin taeniurus Dorsal surface of frontal and parietals of head gray, brown, or olive, with or without a few darker flecks; no apical scale pit present (characters from type description) .........eseeeeseeceeees carajasensis Dorsal surface of frontal and parietal scales of head gray, brownish or live, with a whitish U, V, Xe OG Y mark with exterior black edging, mark may extend onto the internasals; one apical scale pit present MSc Aae rea Sie win Slot Osis. san a caper vacates shaiit tal one Val's) Gj eialereleastorononener's almadensis Dorsal scale LrowsS 19-19-15 2... 2c ccccc ccc cccsceseccccsccccccnccs Dorsal scale rows 19-19-13 ......cccececcerccweccsccce perfuscus Dorsum bright green; head occasionally blue, with or without dark chevrons dorsally and/or laterally porsili Partibie butmevamencctbod satyty. catbex oft .soqtaee em Ventral s <168 pars. ccs) atejegeyete shekeaate steps “tausteted tel Ps quentheri Dorsum with black and/or yellow longitudinal stripes............ Dorsum variable but without stripes.......... cece e cree eceeceees 38 40 41 42 43 46 43. 44. 45. 46. 47. 48. om Lom -*) Dorsum blotched, ground color olive or brown, with darker blotches above and below narrow yellow paravertebral) Mminesees ..)005% Jee NRORES RIE © OU 8 US Le oe Dorsum striped, ground color tan or light brown, with three black stripes, or three black and two yellow stripes, the median stripe beginning on the snout eoeeeeeeeceeeeceeeeeeeoecee ee eee e oe eee eee eee eee eoeoeee ere ee eo cece eee Dorsum with three dark and two narrow yellow stripes, yellow stripes and median dark stripe begin on snout Remini c sie Ses Se see eee ss eee eee. fal ee ee meridionalis Dorsum with three wide black stripes, median black Stripe begins Ono SHOULPeS ee. See LG YO RORe ASL ale . dilepis Ventrals <166; maxillary teeth?13=17 222025 ...20228.8 28 anomalus Ventrals 5176; maxilTaryoteeth?ls-202 24282 20. fe 4 vanzolinii Dorsum uniform or variable, but never as below ................. Dorsum tan or yellowish, with four rows of rounded black, reddish, or reddish brown spots, lateral row of spots smaller than paravertebral row Stele Silo siqioyeld's sas sc Mee MOS eke ees bo oe be sagittifer (see 35) Dorsum-plotched) or with’ squarish Spots’ ..°.......cc.ceec.edse et Dorsum uniform brown to chocolate brown .............. perfuscus Head gray to black, occasionally with light or dark markings, but never red, no middorsal reddish stripe, and generally without rounded black, WNLCHCallany eSPOLSmer’< vot cee ote ote ao «cw ov bales 6 ade Rr www Saree Head black, spotted with red; dorsum with large paravertebral black blotches on olive ground color, rounded lateral black intercalary spots, broad diffuse median ruddy stripe from head to tail, frequently invaded by dorsolateral black spots Beret rensie os eM re ey evar chee, Ootela DONG Grae em dae Sievers wand ance elegantissimus 45 44 47 48 49 49.a Ground color usually gray or tan with a broad whitish dorsolateral line from midbody to tail on scale rows 4, 5 and 6, or 5 and 6;body with small squarish blotches slightly to moderately darker than ground color; venter checkered with black and some shade of pink or red maxillary teeth 19-25 (mode 21-22); diameter of eye 56-78% (x= 66%) of snout length b Ground color usually brown to almost black, one subspecies with an ill-defined, posterior dorsolateral whitish stripe on scale rows 5 and 6. Head usually uniform brown or black; if brown, cephalic scales edged with black; dorsal color pattern highly variable, from salt and pepper pattern to blotches, bands, reticulations or combinations of the above and in many colors; venter from immaculate white to almost black. Maxillary teeth 13-21 (mode 17-18); diameter of eye 41-66% (x= 54%) per cent of snout length ....... poecilogyrus GENUS LIOPHIS Wagler, 1830:187 Liophis almadensis (Wagler) Natrix almadensis Wagler 1824:30. Type locality: Almada, Bahia, Brasil. Natrix almada Wagler 1824:30. (as above) Liophis conirostris Giinther 1854:46. Type locality: Brasil and Bahia. L. (iophis) verecundus Jan 863:300. Type locality: unknown. Liophis wagleri Jan 1863:297. Type locality: Brasil and Bahia. Liophis (Lygophis) y-graecum Peters 1882:129. Type locality: Villa de Guaratinqueta, Sdo Paulo (Brasil). Trigonocephalus scolecomorphus Bacque 1906:116. Type locality: Asuncion, Paraguay. Liophis almadensis Wagler 1830:188. DISTRIBUTION: Chaco Boreal plant formation of Bolivia, all plant formations of Brazil except for Caatinga and Amazon rainforest; Paraguay except for cropland situations in central Paraguay. (published reports of this taxon from the state of Misiones, Argentina, have not been verified). SUBSPECIES: none described; possibly one undescribed cryptic subspecies in Argentina and western Bolivia. Liophis andinus Dixon Liophis andinus Dixon 1983:129. Type locality: Incachaca, 2500 m, Cochabamba, Bolivia. DISTRIBUTION: Known only from the type locality (Figure 2, p. 118, Dixon, 1983). Liophis anomalus (Giinther) Coronella anomala Giinther 1858:37. Type locality: banks of the Rio Parana. Lygophis rutilis Cope 1862:80. Type locality: Rio Tigre, a tributary of the Rio Parana, Paraguay. Coronella pulchella Jan 1863:251. Type locality: Buenos Aires, Argentina. Liophis anomala Amaral 1925:7. DISTRIBUTION: Southern South America, from northwestern Argentina (Salta) east-southeast through Uruguay to Pérto Alegre, Brasil, south to the province of Buenos Aires, Argentina. Liophis atraventer Dixon and Thomas Liophis atraventer Dixon and Thomas 1985:260. Type locality: Boracéia, Sado Paulo, Brasil. DISTRIBUTION: Known only from the type locality (= Estacao Biologica da Boracéia) (figure 4, p. 188, Dixon, 1987). Liophis breviceps Cope Liophis breviceps Cope 1860:252. Type locality: Suriname. Liophis canaima Roze 1957:188. Type locality: Rio Ugueto, Amazonas, Venezuela. DISTRIBUTION: Eastern flanks of the Andes of Ecuador and Peru, eastward to Obidos, Brasil; on the north from central Colombia and northern Guyana south to Rio Mamoré (Trinidad) Bolivia, and Posto Diuarum, Brasil. SUBSPECIES: Two. eae “WeniiaaltS, SUGGS) .s0.¢s,.ccrs ccscitocnya.e .sterelace soe eee breviceps breviceps Be Ventral, Slvr. 1c.. «cle cheyerekers,s spsleres qerspaeseiourene breviceps canaimus Liophis breviceps breviceps Cope 1862:252. Type locality: Suriname. DISTRIBUTION: Forested parts of the Amazon Basin in Bolivia, Brasil, Colombia, Pert, and similar areas in French Guiana, Guyana, Suriname, and Venezuela (see figure 3, p. 154, Dixon 1983). Liophis breviceps canaima Roze 1957:188. (see above) DISTRIBUTION: Known only from the region of the Rio Ugueto, Amazonas, Venezuela (see figure 3, p. 154, Dixon 1983). Liophis carajasensis Cunha, Nascimento and Avila-Pires Liophis carajasensis Cunha, Nascimento and Avila-Pires 1985:53. Type locality: Campo Rupestre d. N 1, Serra Norte, Para, Brasil. DISTRIBUTION: Known only from the type locality (see figure between pp. 18 & 19, Cunha, et al 1985). Liophis cobellus (Linnaeus) Coluber cobella Linnaeus 1758:218. Type locality: America. Coluber serpentinus Daudin 1803:87. Type locality: none given. Coluber cenchrus Daudin 1803:292. Type locality: Asia. Liophis taeniogaster Jan 1863:292. Type locality: Brasil and South America. Liophis trebbaui Roze 1958:262. Type locality: Auyan Tepui, Bolivar, Venezuela. Liophis ingeri Roze 1958:303. Type locality: Chimanta Tepui, Bolivar, Venezuela. DISTRIBUTION: From Villavicencio, Colombia, south to Buenavista, Bolivia, and from Trinidad and Carapito, Venezuela, southeast to Bahia, Brasil. SUBSPECIES: Four. Tea VeEntralliS: S171. nds neniscurere ce sciences se saereeieioee cobellus ingeri BY “Ventiralis,:<1G4) 0); ss). ars cteus, © elerslote o.oieeverroie crete volnenatehatsccps eeaktetetennatsterersarene 2 10 am Ventral, black bands, 2A 5a sr fonas i laveryaistenasbeterdestore fare » cobellus cobellus BRIER tied babliaC ka DANG: <6 hens cecearsede easiest slate sstocldacers os oe o's se sede ve 3.a Ventrals vary from 130-147 (x= 138.4) ......... cobellus dyticus b Ventrals vary from 143-164 (x= 151.6) ..... cobellus taeniogaster Liophis cobellus cobellus (Linnaeus) 1758:292. SYNONYMS: serpentinus Daudin, cenchrus Daudin. DISTRIBUTION: Eastern Guyana Shield, exclusive of Venezuela Tepui system; also Trinidad (see figure 4, p. 158, Dixon 1983). Liophis cobellus dyticus Dixon 1983:159. Type locality: Monte Carmelo (= Requena), Loreto, Peru. DISTRIBUTION: Western Amazon Basin, from Loma Linda, Colombia, south to Buenavista, Bolivia, east to Pérto Velho, Brasil (see figure 4, p. 158, Dixon 1983). Liophis cobellus taeniogaster Jan 1863:292. (see above) DISTRIBUTION: Northeast Brasil, from Isla Bananal, east to central Bahia, north to Rio Amazonas (south bank) (see figure 4, p. 158, Dixon 1983). Liophis cobellus trebbaui Roze 1958:262. SYNONYM: ingeri Roze. DISTRIBUTION: Known only from the Chimanta and Auyan Tepuis, and km marker 144 of the El Dorado-Santa Elena highway, Bolivar, Venezuela (see figure 4, p. 158, Dixon 1983). Liophis cursor (Lacépéde) Coluber cursor Lacépéde 1789:96. Type locality: Martinique. Coluber fugitivus Donndorf 1798:206. Type locality: Martinique. Liophis putnami Cope 1862:78. Type locality: Martinique. Liophis cursor andreoides Werner 1924:36. Type locality: Grenada. DISTRIBUTION: Known only from the island of Martinique, West Indies (see figure 1, p. 297, Dixon 1981). Liophis dilepis (Cope) Lygophis dilepis Cope 1862:348. Type locality: Paraguay. Liophis dilepis, Dixon 1980:7. DISTRIBUTION: From northeast Brasil, south to southern Brasil, and Paraguay, extending northwest to southern Bolivia (see figure 1, p. 4, Michaud and Dixon 1987). 11 Liophis elegantissimus (Koslowsky) Rhadinaea elegantissima Koslowsky 1895:115. Type locality: Sierra de la Ventana, Province of Buenos Aires, Argentina. Liophis elegantissimus, Amaral 1925. DISTRIBUTION: Known from five localities within the Sierra de la Ventana, Buenos Aires, Argentina (see figure 1, p. 566, Dixon 1985). Liophis epinephelus Cope Liophis epinephelus Cope 1862:78. Type locality: Truando, Colombia. Liophis reqinae albiventris Jan 1863:294. Type locality: Western Andes of Ecuador (fra Lacutunga e Guayaquil) (= Latacunga?). Liophis reginae quadrilineata Jan 1863:295. Type locality: Ecuador. Ophimorphus alticolus Cope 1868:102. Type locality: Quito Valley, Ecuador. Zamensis ater Giinther 1872:22. Type locality: Biscra, Algeria (in error). Liophis fraseri Boulenger 1894:131. Type locality: Western Ecuador. Liophis bimaculatus Cope 1899:71. Type locality: Colombia. Liophis bipraeocularis Boulenger 1903:351. Type locality: Facatativa, Colombia. Liophis opisthotaenia Boulenger 1908:114. Type locality: Mérida, Venezuela. Liophis pseudocobella Peracca 1914:99. Type locality: Angeldpolis, Colombia. Liophis cobella alticolus Amaral 1931:87. Type locality: Jericé, Colombia. Liophis taeniurus juvenalis Dunn 1937:213. Type locality: San Jose, Costa Rica. Liophis bimaculatus lamonae Dunn 1944:486. Type locality: Sonsén, Antioquia, Colombia. Liophis epinephelus ecuadorensis Laurent 1949:8. Type locality: Ecuador. DISTRIBUTION: Trans-Andean South America from Venezuela to Peru; Cis-Andean South America at and above 2,200 m in Ecuador and Pert; also Central America from central Costa Rica through Panama (see figures 1, 2, and 3, pp. 133-135, Dixon 1983). SUBSPECIES: Eight. l.a Lateral black tail stripe absent, dorsal and ventral surfaces of body and tail alternately marked with red, and black 2. of. LPS oe . Pode cheted rere s COR es oo Fe SNOT Cnet 2 b Lateral black tail stripe present, dorsum may be banded, spotted, flecked, or almost unicolor, belly checkered with«black. or notaas...iseo.tece bites. Pbsmeraeeseer ss eee ose es. 3 2:a Subcalidailis <547.. 5 ..c5.d0da0 se aetem serene epinephelus pseudocobellus breSubcaudall’s; 5540. 32 .ntw ance... Gotaapeo.. See epinephelus juvenalis 32a” Ventyalls <165) 2.5. ais cecaicictave evarshe oitaccve iia ieregel cue har enrertetetarrerenhets lehetenete 4 bePVenthal'SwSll65) .. sy. eis oc whajetore s 76 or more, juveniles with a pair of light nuchal spots, adults uniform brown, each dorsal scale witha pale Vight center?s..3 200.5655. .55;3 miliaris amazonicus b Subcaudals <68; juvenile pattern variable, but never with a pair of light nuchal spots, adults light tan to black, with light centered scales, or with light flecks, Scattered. EhYOUGHOUES © crciis ] 2 '« sagittifer modestus b Dorsum with large to median sized paravertebral reddish black to black blotches from nape to tail; often with secondary row of intercalary black blotches laterally, and occasionally a median series of dorsal blotches anteriorly; 100% of population with 19-19-15 scale OWS) Sellar a; SA at eee Pe ecco che sagittifer sagittifer Liophis sagittifer sagittifer (Jan) 1863. SYNONYMS: pulcher Steindachner; argentinus Bréthés. DISTRIBUTION: Monte of Patagonia, from Tucuman, south to the state of Chubut, Argentina. (see figure 2, p. 391, Dixon and Thomas 1982). : Liophis sagittifer modestus (Koslowsky) 1896. SYNONYM: trifasciatus Werner. 23 DISTRIBUTION: Chaco-Bonariesian Plain of Argentina, Bolivia, and Paraguay. (see figure 2, p. 391, Dixon and Thomas 1982). Liophis taeniurus Tschudi Liophis taeniurus Tschudi 1845:164. Type locality: Peru, in der heissen waldregion. DISTRIBUTION: Middle and upper elevations of the Andes in southern Ecuador, south through Pert to the Cochabamba region of Bolivia. SUBSPECIES: None described; possibly two or more suggested from recent analysis of data from additional material. Liophis triscalis (Linnaeus) Coluber triscalis Linnaeus 1758:224. Type locality: India (in error). Coluber corallinus Linnaeus 1758:223. Type locality: Asia (in error). Liophis triscalis, Boulenger 1894:129. DISTRIBUTION: Known only from the Leeward Island of Curacao. (see figure 1, p; 2975.Dixon) 1981): Liophis typhlus (Linnaeus) Coluber typhlus Linnaeus 1758:218. Type locality: India (in error). Xenodon isolepis Cope 1870:155. Type locality: Pebas, Ecuador (= Perd) Opheomorphus brachyurus Cope 1887:57. Type locality: Chupada, Mato Grosso, Brasil. Liophis elaeoides Griffin 1916:187. Type locality: Prov. del Sara, Bolivia. Liophis macrops Werner 1925:57. Type locality: Paramaribo, Suriname. Liophis typhlus, Dixon 1980:16. DISTRIBUTION: Rainforests of the Guiana Shield and Amazon Basin, also the Chaco and Cerrado of Bolivia, Brasil, and Paraguay. (see figure 1, p. 175, Dixon 1987). SUBSPECIES: Three. l.a_ Ventrals 133-163 (x= 147.3); juveniles and adults with dark paravertebral chevron marks ........ typhlus typhlus b Ventrals 158-172 (x= 163.5); juveniles and adults without? dark ichevnon iMaricSiee ru. cteith at tatalciens aeetebotattode choplcteush teueve ote. os 2.a Subcaudals 40-49 (x= 44.4); tail/total length ratios Os 1A O-OR1GO? (R= Or 4O seer tie steRiet cece typhlus brachyurus b Subcaudals 49-56 (x= 52.0); tail/total length ratios ONGO= 04200) m=O? Us) rieate 6 ap2'cs eiecs aser ors overseers typhlus elaeoides Liophis typhlus typhlus (Linnaeus) 1758. SYNONYMS: isolepis Cope; macrops Werner. 24 DISTRIBUTION: Rainforests of Guiana Shield, and Amazon Basin (see figure ip. 175, Dixon 1987). Liophis typhlus brachyurus (Cope) 1887. DISTRIBUTION: Deciduous mesophytic forests of southeastern Brasil, and the Campo Cerrado forests of east-central Brasil (see figure 1, p. 175, Dixon 1987). Liophis typhlus elaeoides Griffin 1916. DISTRIBUTION: Mesic Chaco forests of southeastern Bolivia; northern Paraguay, and western Mato Grosso, Brasil (see figure 1, p. 175, Dixon 1987). Liophis vanzolinii Dixon Liophis vanzolinii Dixon 1985:567. Type locality: Achiras, Cordoba, Argentina. DISTRIBUTION: Known only from three localities in the western part of the Argentine state of Cordoba (see figure 1, p. 566, Dixon 1985). Liophis viridis Giinther Liophis viridis Giinther 1862:58. Type locality: Pernambuco, Brasil Liophis typhlus prasina Jan and Sordelli 1866:18(4)(3). Type locality: Brasil. DISTRIBUTION: The Caatinga, Agreste, and Atlantic rainforests of Brasil. (see figure 2, p. 181, Dixon 1987). SUBSPECIES: Two. l.a Ventrals 169-188 (x= 177); reduction site ventrals BSA UG (SSR OG GY) nt Se cu wed ova ns LAS IS. desl aks oe viridis viridis b Ventrals 181-202 (x= 189.8); reduction site ventrals MOP ai 25 tele mmeLNA) Gi) Besa, «015.01, cvsnne. oie dal obs alet Tahsh bake ds viridis prasinus Liophis viridis viridis Giinther 1862. DISTRIBUTION: The agreste and Atlantic rainforests of Brasil, from Recife to Salvador (see figure 2, p. 181, Dixon 1987). Liophis viridis praesinus Jan and Sordelli 1866. DISTRIBUTION: The Caatinga forest of Brasil (see figure 2, p. 181, Dixon 1987). Liophis williamsi (Roze) Urotheca williamsi Roze 1958:1. Type locality: E1 Junquito, D.F., Venezuela. Liophis williamsi, Dixon 1980:17. 25 DISTRIBUTION: Cloud Forests of the coastal Andes of Venezuela (see figure 2, p. 118, Dixon 1983). 26 COUNTRY LISTS (containing taxa of Liophis) CENTRAL AMERICA BRASIL COSTA RICA Liophis atraventer Liophis almadensis Liophis epinephelus juvenalis Liophis anomalus Liophis breviceps breviceps PANAMA Liophis carajasensis Liophis cobellus dyticus Liophis epinephelus juvenalis Liophis cobellus taeniogaster Liophis epinephelus epinephelus Liophis dilepis Liophis lineatus Liophis flavifrenatus Liophis frenatus SOUTH AMERICA Liophis jaegeri jaegeri Liophis jaegeri coralliventris ARGENTINA Liophis lineatus Liophis longiventris Liophis sp. nov. Liophis maryellenae Liophis anomalus Liophis meridionalis Liophis elegantissimus Liophis miliaris amazonicus Liophis flavifrenatus Liophis miliaris chrysostomus Liophis quentheri Liophis miliaris merremii Liophis jaegeri coralliventris Liophis miliaris mossoroensis Liophis miliaris semiaureus Liophis miliaris orinus Liophis meridionalis Liophis miliaris semiaureus Liophis poecilogyrus Liophis paucidens Liophis reginae macrosomus Liophis poecilogyrus Liophis sagittifer modestus Liophis reginae macrosomus Liophis sagittifer sagittifer Liophis reginae semilineatus Liophis vanzolinii Liophis typhlus brachyurus Liophis typhlus elaeoides BOLIVIA Liophis typhlus typhlus Liophis viridis prasinus Liophis almadensis sub-sp. Liophis viridis viridis Liophis almadensis almadensis Liophis andinus Liophis breviceps breviceps COLOMBIA Liophis cobella dyticus Liophis dilepis Liophis breviceps breviceps Liophis quentheri Liophis cobellus dyticus Liophis meridionalis Liophis epinephelus bimaculatus Liophis miliaris amazonicus Liophis epinephelus epinephelus Liophis poecilogyrus Liophis epinephelus lamonae Liophis reginae macrosomus Liophis epinephelus opisthotaenius Liophis reginae semilineatus Liophis epinephelus pseudocobel lus Liophis sagittifer modestus Liophis festae Liophis taeniurus Liophis lineatus Liophis typhlus elaeoides Liophis melanotus melanotus Liophis typhlus typhlus Liophis miliaris chrysostomus Liophis reginae semilineatus Liophis typhlus typhlus ECUADOR Liophis breviceps breviceps Liophis cobellus dyticus Liophis epinephelus albiventris Liophis epinephelus bimaculatus Liophis epinephelus epinephelus Liophis epinephelus fraseri Liophis epinephelus ]amonae Liophis festae Liophis lineatus Liophis miliaris chrysostomus Liophis reginae semilineatus Liophis taeniurus Liophis typhlus typhlus FRENCH GUIANA Liophis breviceps breviceps Liophis cobellus cobellus Liophis lineatus Liophis miliaris miliaris Liophis poecilogyrus Liophis reginae reqinae Liophis typhlus typhlus GUYANA Liophis breviceps breviceps Liophis cobellus cobellus Liophis lineatus Liophis miliaris miliaris Liophis poecilogyrus Liophis reginae reqinae Liophis typhlus typhlus PARAGUAY Liophis almadensis Liophis dilepis Liophis flavifrenatus Liophis frenatus Liophis quentheri Liophis jaegeri coralliventris Liophis longiventris Liophis meridionalis Liophis miliaris semiaureus Liophis poecilogyrus Liophis reqinae macrosomus Liophis sagittifer modestus Liophis typhlus elaeoides 27 PERU Liophis breviceps breviceps Liophis cobellus dyticus Liophis epinephelus fraseri Liophis festae Liophis miliaris chrysostomus Liophis problematicus Liophis miliaris chrysostomus Liophis reginae semilineatus Liophis taeniurus Liophis typhlus typhlus SURINAME Liophis breviceps breviceps Liophis cobellus cobellus Liophis lineatus Liophis miliaris miliaris Liophis poecilogyrus Liophis reginae reginae Liophis typhlus typhlus URUGUAY Liophis anomalus Liophis jaegeri jaegeri Liophis miliaris semiaureus Liophis poecilogyrus VENEZUELA Liophis breviceps breviceps Liophis breviceps canaimus Liophis cobellus cobellus Liophis cobellus trebbaui Liophis epinephelus opisthotaenius Liophis lineatus Liophis melanotus melanotus Liophis melanotus subspecies Liophis poecilogyrus (?) Liophis reginae semilineatus Liophis reginae zweifeli Liophis typhlus typhlus Liophis williamsi CARIBBEAN ISLANDS BARBADOS Liophis perfuscus 28 CARIBBEAN ISLANDS (Continued) CURACAO Liophis triscalis DOMINICA Liophis juliae GUADELOUPE Liophis juliae MARIE-GALANTE Liophis juliae MARTINIQUE Liophis cursor SAINT LUCIA Liophis ornatus TOBAGO Liophis melanotus Liophis reginae TRINIDAD Liophis cobellus cobellus Liophis melanotus subsp. Liophis reginae zweifeli SOUTH AMERICA . festae - guentheri / . viridis - jaegeri SCALE 300 400 500 600miLES KILOMETERS SOUTH AMERICA No. 3 lineatus | meridionalis | maryellenae i atraventer SCALE 100 200 300 400 Soo 600mILES 200 400 00 800 KILOMETERS SINUSOIDAL PROJECTION 30 WEST LONGITUDE GOODE BASE MAP SERIES DEPARTMENT OF GEOGRAPHY THE UNIVERSITY OF CHICAGO WGNRY A LEPPARD, EDITOR SUUTH AMEKICA s No. 3 [=] L. witliamsi | L. almadensis | 500 600 MILES 00 600 KILOMETERS SINUSOIDAL PROJECTION / \ 30 WEST LONGITUDE GOODE BASE MAP SERIES Propared by Haney M. Leppard DEPARTMENT OF GEOGRAPHY FR ee ee ae CHICAGO SOUTH AMERICA epinephelus : | reginae / problematicus | andinus 500 600 MILES 400 600 800 KILOMETERS SINUSOIDAL PROJECTION GOODE BASE MAP SERIES GEOGRAPHY OGPARTMENT OF THE WEST LONGITUDE SOUTH AMERICA Ars L. anomalus (*] L. vanzolinii / [=| Ee. elegantissimus | 500 600 mMLES 600 600 KILOMETERS SINUSOIDAL PROJECTION GOODE BASE MAP SERIES GEOGRAPHY DEPARTMENT OF THE UNIVERSITY WEST LONGITUDE SOUTH AMERICA - No. 3 Jal L. dilepis 4 | ; L. paucidens SCALE 100 200 300 400 500 600MmiLES 200 400 600 800 KILOMETERS SINUSOIDAL PROJECTION 30 WEST LONGITUDE GOODE BASE MAP SERIES Pryce by Hew M. Lopord DEPARTMENT OF GEOGRAPHY © ret be Te — THE UNIVERSITY OF CHICAGO. HENRY M LEPPARD, EDITOR SOUTH AMERICA No. 3 ° ! ° e 500 600 MILES 800 KILOMETERS SINUSOIDAL PROJECTION ‘i SS SOUTH AMERICA ; ee 80 pose De eT INTPAU th Ni mh Troe ie Ee ag ‘a ; SSESNa Saas i= ae = : “ gk als Hea sear Y < hy BA melanotus / L. miliaris SOUTH AMERICA : No. 3 SCALE 100 200 300 400 S00 s600MmiLES 600 MILOMETERS SINUSOIDAL PROJECTION / SOUTH AMERICA WS py 7A , JN) L. breviceps | | Es longiventris SCALE ° 100 200 300 400 500 600MmMILES 200 400 00 800 KILOMETERS SINUSOIDAL PROJECTION 30 WEST LONGITUDE SOUTH AMERICA ’ Ne 3 -sagittifer | typhlus 600 MILES 800 KILOMETERS SINUSOIDAL PROJECTION 30 WEST LONGITUDE Montserrat a} L. juliae L. triscalis A Curacao EAN > VENEZUELA Guadeloupe Oe. Desirade - Marie-Galante oe Dominica L. cursor — > Martinique L. ornatus ee ge Barbados (OS Gir San Vicente a bo, GO’ The Grenadines + p Grenada ie cobellus Trinidad L. melanotus SS 1 ( :. b }' Se ee Ph slovs bse? : of a = rks * 7g —h ee ; ejneal¥ a a ee Aa! ; ‘> ’ —. ‘a oe ‘a eonivanera ae a 4, ST =e | | Seedioe) oJ auitedos at eae a6 pabintit } i at - . 9(rulleds tan tincorrect Original spelling, non-existent nomenclaturally, of Sphenodon Gray, 1831, type species, Hatteria punctata Gray, 1842; emended first to Sphenodon by Gray, 1872, and thus accepted by the EC ZAIN ins Sphalerosophis Jan, 1865, in Filippi, Note Viaggio Persia:356 (an incorrect original spelling and jr. synonym of Spalerosophis Jan, 1865; Sphalerosophis Jan is therefore non-existent nomenclaturally). Spiarquiss Merrem,. Uuec0),.) Tente syst. sAmph. 219 iar jm. =synonymaot Dermochelys Blainville, 1816, having the same type species, resiudecomvacem. Vandeldu,*°1761 7) in= the case” of Sphargis, Ehzough a 7m. Synonym of To coriacea, Ss. mercurialis~Merrem, IL tS32(0))) Stegoporus Wiegmann, 1832, in Wiegmann and Ruthe, Handb. Zool., Ed. 2:204 (rejected in priority but not homonymy contexts in Onder eOm protect Ambystzoma’ Tsichuday,.el8s8; proposed= as a substitute for Siredon Wagler, 1830, a jr. synonym of Gyrinus mexicanus Shaw, 1898, now in Ambystoma). Ste Lio, Daudi ns 802,.HisSt.wNat.. ~Rept...» 43.5: ola. .j%..-homonyms of Stellio Laurenti, 1768, a nomen dubium because its type species, S. saxatilis, is unidentifiable; neither Laurenti name has been rejected officially, however, and Stellio Laurenti has commonly been regarded as an invalid sr. synonym of Agama Daudin, 1802). Tachyophis Mertens, 1934, Arch. Naturg. (n.f.) 3:197 (invalid as a jx. homonym of Tachyophis Rochebrune, 1884, a fossil snake; type species Coluber pictus Gmelin, 1788, now placed in Dendrelaphis Boulenger, 1890, as a valid species). TaparavOken,; | 18167, lehrb. Natur. 3:vi9 (index)? (this “work “by Oken has been rejected for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally, as of Oken, 1816, and should not be cited in the synonymy of anything, although commonly referred to Phrynosoma Wiegmann, 1828; also an incorrect original spelling of Tapaia Oken, 1816, q. v.). 2 Tapaia Oken, 2817; sIsas von) Oken 911817: 1183) ejected sinispritoxalty, but not in homonymy contexts to protect Phrynosoma Wiegmann, 1828, having the same type species, Lacerta orbicularis Linnaeus, 1758). Tapaja Oken, 1816, Lehrb. Naturg. 3:295 (this work by Oken was rejected for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of anything). Tapaya Fitzinger, 1826, Neue Classif. Rept.:17 (rejected in priority but not homonymy contexts to protect Phrynosoma Wiegmann, 1828, having the same type species, Lacerta Orbiicularius! Linnaeus), 147/518)).. Tapaya Oken, 1817, Isis von Oken 1817:1183 (an’incorrect original s\peldbinige Mode vapiativg Oken, iey Wa hence non-existent nomenclaturally). LapaviasGray,,)1825, sAnn. Philos. 26:1 975 (rejected. in, priority but not in synonymy contexts to protect Phrynosoma Wiegmann, 1828, having the same type species, Lacerta orbicularis Linnaeus, 17:98 ); z Thermophilus Fitzinger, 1843, Syst. Rept.:21 (rejected in priority but not homonymy contexts to protect Ichnotropis Peters, 1854, both having the same type species, Algyra capensis A. Smith, 1838, in Thermophilus via Fitzinger's Tropidosaura capensis Dune rnis sands. Babron ws anime Gores Ons Aww omic hy), sand ein, alichnotinopss via I. macrolepidota Peters, 1854, a jr. synonym of A. capensis Smith) 2 Ope 4228 Leet Oppel) sis “GAnne! UiMusareNat® . He Si GaNicts ekarus 16(95) :377,381 (rejected as a jr. homonym of Tortrix Linnaeus, 1758, a lepidopteran insect; type species Anguis scytale Linnaeus, 1758, still valid, now referred as type species to the genus "Apis Oken,;niel6, of whitch lortrige Oppel as) dics. synonym, but invalid). Lritropis Fitzinges,, 1843, Syst. Rept. :59 (rejected in, priority but not in homonymy contexts to protect Chalarodon Peters, 1854, having the same type species, Tropidogaster blainvillii Duméril and Bibron, 1837). Tropidogaster Duméril and Bibron, 1837, Erp. Gen. 4:329 (rejected in priority but not in homonymy contexts to protect Chalarodon Peters, 1854, having synonymous type species; see blainvillii in the species-group list) 3 Typhlina Wagler, 1830, Nat. Syst. Amph.:196 (rejected in priority bute not Han shomonymy, fcontextis «tos protect leptotyphilops Fitzinger, 1843, type species Typhlops nigricans Schlegel, 1839, in which genus Anguis septemstriatus Schneider, 1801, the type species of Typhlina, also belongs). Zygnis Oken, 1816, Lehrb. Naturg. 3:284 (this work by Oken has been rejected for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of anything). SPECIES-GROUP NAMES alamose, Kinosternon, Pritchard, 1979, Encyclopedia of Turtles: 556 (rejected in both priority and homonymy contexts to protect K. alamosae Berry and Legler, 1980; a non-existent name, nomenclaturally, that should not be cited as a synonym of anything). alleghaniensis, Abranchus, Harlan, 1825, Ann. Lyceum Nat. Hist. New York 1(18):271 (an unjustified emendation and jr. synonym of Salamandra alleganiensis Daudin, 1803, now Cryptobranchus alleganiensis (Daudin)). aulilucgaiviow, bacermta,y. Blumenbach, | 17/719), Handb. Natuxrg:. (1:2'63 (rejected in priority but not in homonymy contexts to protect Spaced sins Missussippiensas Daudin,;,1801,, now Alligator ississippiensis (Daudin), of which it is a jr. synonym). arnreollatia, shacerta, sHouttuyn,.wWiei >. Anwme Mus ..Houtt.. Index: 24 (this work by Houttuyn was suppressed for nomenclatural purposes (Opinion 380), hence this name should not be cited in the synonymy of anything, as it does not exist nomenclaturally, and is in addition unidentifiable). arias scolubey. Gmelin, I786-s rm Lannaeus, Syst. Nat., sa. 3) 1: 1103 (rejected in both priority and homonymy contexts, by Gmedin forisany other sauthor, “prior, to erection: of »Coluber atratus Hallowell, 1845, now Ninia atrata (Hallowell), in order to protect the latter name; previous usages do not exist nomenclaturally and should not be cited in the synonymy of anything). besseri, Anguis, Andrzejowski, 1832, Nouv. Mém. Soc. Imp. Nat. Moscoul (2) 2-330. stab.) 22 tig... 7,, tab. 24. (rejected in priority but not in nomena contexts to protect QOtophis eryx colchica Notdmann, 1840, now Anauis fragilis colchi'ca, “of which A. bessemigits Now a sii Synonym)! . 14 bibronii, Trapelus (Psammorrhoa), Fitzinger, 1843, Syst. Rept. 81 (rejected in both priority and homonymy contexts to eeeae Agama bibronii A. Duméril in Duméril and Bibron, 1851; Fitzinger's name was a sr. secondary homonym of Duméril's, having been transferred to the synonymy of Agama hispida aculeata Merrem, 1820; it does not now exist nomenclaturally). bilineatus, Psammophis moniliger, Peters, 1867, Monatsb. Akad. Wiss. Berlin 1867: 237 S(seyected wan Sprilomilsy shut inet an homonymy contexts to protect P,. sibilans subtaeniata Peters, 1882, now PB. subtaeniata Peters, of which bilineatus is now a jr. synonym). blainvillii, Tropidogaster, Duméril and Bibron, 1837, Erp. Gen. 4:300 (rejected in priority but not in homonymy contexts to protect Chalarodon madagascariensis Peters, 1854, of which blainvillii is now a jr. synonym). boseiRand, Bory cde St. Vincent, 1828, Res. Erp. : 266 (rejected in priority but not in homonymy contexts, to protect Rana esculenta perezi Seoane, 1885, now Rana perezi Seoane, of which the former is now a jr. synonym). caesius,, Coluber, Cloquet, 1818, Dilers Sci. “Nata 201 (rejected in priority but not in homonymy contexts to protect Coluber irregularis Leach, 1819, now Philothamnus irregularis (Leach), of which the former is now a jr. synonym). californiana, Aspidonectes, Rivers, 1889, Proc. California Acad. Soi.. (2)' 22) 233-236 (rejected in prionity but mot in homonymy. contexts to protect Trionyx steindachneri Siebenrock, 1906, of which the former is now a jr. synonym). Chiametla, Coluber, Shaw, 1802, Gen. Zool. 3(2): 440 (rejected in priority but not in homonymy contexts to protect Herpetodryas Margaritiferus Schlegel, 1838, now Drymobius margaritiferus (Schlegel), and Drymobius margaritiferus fistulosus Smith, 1942, of which Shaw's name is now a jr. synonym) . cincolor, Crotalus durissus, Notestein, 1905, 7th Rep. Michigan INGO IE WIS GALS 2 123 (ruled non-existent nomenclaturally because cited -only, in, synonymy, of Crotalus, hopridus Linneeus, . 1758; presumably a lapsus for concolor, and presumably drawn from Jan, 1859, although the only source stated was "J"). cinereous,.Crotalus, Le, Conte,. 19:52)! tiny Ha blowelil,s/proc. \Acad: Nat. S¢1., Philadelphia 5,(5).:, 177, .(mejected, an. priority. but.~not in homonymy contexts to protect Crotalus atrox Baird and Girard, 1853, of which the former in now a jr. synonym). tS coerulea, Rana, Houttuyn, 1787, Anim. Mus. Houtt. Index: OA GEhaS work by Houttuyn has been suppressed for nomenclatural purposes (Opinion 380), hence the name does not exist nomenclaturally and is unidentifiable anyhow). colonorum, Agama; ~Daudin, 1802, Hist.qNat.-Rept< 3: SSio7 (ay air objective synonym of Lacerta agama Linnaeus, 1758, now A. a. agama (Linnaeus)). concolor, Crotalus durissus, Garman, 1883, Mem. Mus. Comp. Zool. 8: 175 (ruled non-existent nomenclaturally because cited only in synonymy, of C. horridus Linnaeus, 1758; name attributed to Vane LS59))e concolor, Crotalus durissus, Gloyd, 1940, Spec. Publ. Chicago Acad. Sci. 4:171 (ruled non-existent nomenclaturally because cited only in synonymy, of C. viridis decolor Klauber, 1930; name attributed to Jan, 1859). concolor, Crotalus durissus, Jan, 1859, Rev. Mag. Zool. (2)10: 153 (ruled non-existent nomenclaturally because a nomen nudum). Gruciger Bufo,” Oken;,) (18i6,;oLehzbin Naturg.. 312209 (thismwork by Oken has been suppressed for nomenclatural purposes (Opinion 417), hence the name is non-existent nomenclaturally and should not be cited in the synonymy of any species). eupreus, Coluber, Houttuyn, 27/87, Anima Must ~Houtt .» index: 28 (this work by Houttuyn has been suppressed for nomenclatural purposes (Opinion 380), hence the name does not exist nomenclaturally and is unidentifiable). digjigis Sits iChiiretes) Saenz po1869),; Ani sUnivewNacwrEStescmidos Colombia 1869: 63 (rejected in priority but not in homonymy contexts to protect Heteroclonium bicolor Cope, 1896, now Bachia bicolor (Cope), of which Saenz' name is now a jr. synonym). Op. 1482. deltiatus ss, Coluber, iinnacuswe, 1766;.9Syste: Nat efznBdvml2ancli: 376 (rejected in priority but not in homonymy contexts to protect Coluber coccineus Blumenbach, 1788, now Cemophora coccinea (Blumenbach), of which doliatus is now a jr. synonym). dorsata, Testudo, Schoepff, 1801, Naturg. Schildk.: 158 (rejected in priority but not in homonymy contexts to protect Testudo Ppunctularia Daudin, 1802, now Rhinoclemmys punctularia (Daudin), of which dorsata is now a jr. synonym). Gracaena, Lacerta, Linnaeus, 1766, Syst. Nat., Ed. 12, 1: 250 (rejected in priority but not in homonymy contexts to protect 16 Tupinambis bengalensis Daudin, 1802, now Yaranus bengalensis (Daudin), of which dracaena is now a jr. synonym). dubia, Amphisbaena, Rathke, 1863, Abh. K.-Bayer. Akad. Wiss. Miinchen 9(1): 128 (rejected in both priority and homonymy contexts to protect Amphisbaena dubia Mliller, 1924; Rathke's name is non-existent nomenclaturally but was based on A. fuliginosa Linnaeus, 1758, and, more precisely, A. £. amazonica Vanizolsmnry. u9iSb)ie: elaphis, Coluber, Shaw, 1802, Gen. Zool. 3: 450 (rejected in priority but not in homonymy contexts to protect Coluber Scalaras Schinz, 1822, now Bilaphe scalaris (Schinz)=!) oftfiwhich Shaw's name in now a jr. synonym). erengicauiday, sLiaittiony mir sichisicholitziam GSsisrir Zools® “Atlas ist 14 (rejected in priority but not in homonymy contexts to protect Salamandra lugubris Hallowell, 1849, now Aneides lugubris (Hallowell), of which Eschscholtz' name is now a jr. synonym). emnyitshiiomiotal, = Siaslamamdicad 7 eRak inesiqueyae 1S WSyeiSica pas tdi aE: 215 (rejected in priority but not in homonymy contexts to protect Salamandra cinerea Green, 1818, now Plethodon cinereus (Green), of which Rafinesque's name is now a jr. synonym) . fasciata, Lacerta, Houttuyn, 1787, Anim. Mus. Houtt. Index: 24 (this work by Houttuyn has been rejected for nomenclatural purposes (Opinion 380), hence the name does not exist nomenclaturally, and besides is unidentifiable). fasciata, Rana, Burchell, 1824, Travels Interior South Africa 2: 3:2, e(rejected..in |iprierity, but..notsjin Hhomonymy ‘contexts to protect Rana grayi Smith, 1849, of which Burchell's name is now a jr. synonym; in addition, all other uses of Rana fasciata prior to that of Smith, 1849, are similarly rejected). flava, Testudo, Dg 1788, Hist. Nat. Quad: Ovap. sSexpenisy a, Synops. Meth. 135,, tab, 16 (rejected san ypriority but not in homonymy contexts to protect Cistudo blandingii Holbrook, 1838, now Emydoidea blandingii (Holbrook), of which Lacépéde's name is now a jr. synonym). flavescens, Amphisbaena, Houttuyn, 1787, Anim. Mus. Houtt. Index: 29 (this work by Houttuyn has been suppressed for nomenclatural purposes (Opinion 380), hence the name does not exist nomenclaturally; it is also unidentifiable). foetidus,, Coluber, Guldenstedt,., 1801, vim ~Georgs,,. Geogr. -Phy si. Naturh. Beschreib. Russ. Reich. 3(7): 1884 (rejected in priority but not in homonymy contexts to protect Pelias renardi 17 Christoph, 1861, now Vipera ursinii renardi, of which foetidus is now a jr. synonym). formosissimus, Constrictor, Laurenti, 1768, Specimen Medicum...: 107 (a jr. objective synonym of Boa constrictor Linnaeus, WTS). funebris)” Salamandra, "Bory de St °-Vincent;, "828; Res. Exrp..:: 2316 (rejected in priority but not in homonymy contexts to protect Pleurodeles walt] Michahelles, 1830, of which funebris is now a jr. synonym). galliwasp, Scincus, Oken, 1816, Lehrb. Naturg. 3: 299(this work by Oken has been suppressed for nomenclatural purposes (Opinion 417), hence the name is non-existent nomenclaturally and should not be cited in the synonymy of any species). Cpaecrs ,LJSied JG) | Oken .hUsr671) Lelicb..aNiat weg.) 3% 202(as in the preceding account of galliwasp) granulatus}; Anguis, Houttuyn, 1787, Anim. Mus. Houtt. Index: ~29 (this work by Houttuyn has been suppressed for nomenclatural purposes (Opinion 380), hence the name does not exist nomenclaturally; it is also unidentifiable). grisea, Lacerta, Oken, 1816, Lehrb. Naturg. 3: 303 (this work by Oken has been suppressed for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally and should not be cited in the synonymy of any species). indicus, Crocodi lus wulgaris, (Gray, 1831),/ Syn., (Rept..2 058 <(regected in priority but not in homonymy contexts to protect Crocodilus palustris Lesson, 1831, now Crocodylus palustris (Lesson), of which Gray's name iS now a jr. synonym). istcUMICWUS Ee StClMeKOne Oken),, SiO, lehbe. Nalteumg. S34 204 (this work by Oken has been suppressed for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of any species). JaAGAapaisa,; Colubes,. Houttuyn,. 17/87... Anim: -Muis,.. Houtt index: 26 (this work by Houttuyn has been suppressed for nomenclatural purposes (Opinion 380), hence the name does not exist nomenclaturally: it is also unidentifiable). lancifer, Trigonocephalus, Oken, 1816, Lehrb. Naturg. 3: 270 (this work by Oken has been suppressed for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of any species). 18 MAMMAL (COMMA Uacacews, Iss. ShySies | Wehe., iwell 10), Weve (rejected in priority but not in homonymy contexts to protect Coluber oceipitomacwiketus, Storer, Lssoetnowes Stioger aa occipitomaculata (Storer), of which it is now a jr. synonym). lepidopus, Bipes, Oken, 1816, Lehrb. Naturg. 3: 249 (this work by Oken has been suppressed for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally and should not be cited as a synonym of any species). Lucius,» Crocodilus (Aldigator), Cuvier; 807, Anns Muse aNatawnists. Nate Panise AsOk: 28) S(a a) Go ees Vy noOnym! Of se CaaxolCloidemlauES mMississippiensis Daudin, 1801, now Alligator mississippiensis (Daudin)). lutescens, Triturus, Rafinesque-Schmaltz, 1832, Atlantic J. Friend of Knowledge 1: 121 (rejected in priority but not in homonymy Contextspito protect sGyrinepha Ins! pomp hwacssit icuis: duasyi) ywWelslexy, 1930, o£ which it is*now a jEqesynonym) = Marmorata, Amphisbaena, Houttuyn, 1787, Anim. Mus. Houtt. Index: 30 (this work by Houttuyn has been suppressed for nomenclatural purposes» ,(Opinsion= 380) .- hence, the » name) sdoes, not sexist nomenclaturally and should not be cited in the synonymy of any species; it is also unidentifiable). maxima, Rana, Laurenti, 1768, Specimen Medicum...: Sy (el sre Synonym, | Of sRanmasboanis) ~binnaeus,;& I S8),_ “noOws ~Hyla ~ bioia nis (Linnaeus)). melanepis, Coluber, Rafinesque-Schmaltz, 1814, Précis Découv. Trav. Semiolog. 15 (rejected in priority but not in homonymy contexts to protect Coluber viridiflavus carbonarius Bonnaparte, 1833, now Hemorrhois viridiflava carbonaria (Bonaparte), of which melanepis is now a jr. synonym). melanocercus, Drymarchon Baws, | Smith, eel 4iy, wis Washi ngtonwAcad: Scns AST A484 (alague tobslectuivjemisy nonymeofe ySipnelrotzets: melanurus Duméril, Bibron and Duméril, 1854, now Drymarchon corais melanurus (Duméril, Bibron and Duméril); originally proposed as a substitute for the latter name, supposed to be invalidated by its senior secondary homonym, Coluber melanurus Schlegel, 1837, via Spilotes melanurus (Schlegel) Gray, 1858). meleagris, “Testudo, Shaw, (1793, "Nate Mise. :78tab.9 44° (rejecteds in priority "but Not “in” Hononymy contexts to protect“ Grstudo blandingii Holbrook, 1838, now Emydoidea blandingii (Holbrook), of which Shaw's name is now a jr. synonym). iL) Merenre dis oO Spheres, iMexcemys Le20AaTent -ASyst...Amph 3:19 (a jr: objective synonym of Testudo coriacea Vandelli, 1761, now Dermochelys coriacea (Vandelli)). machahedllesaavy Podancis, —Fitzinges,;) 1864 =un Erber; Verh. Zool -— Bot. Ges. Wien, 14: 703 eee sp) Jove slopentients #lojbhieas” sovoneis mala! homonymy contexts to protect Lacerta viridis trilineata Bedriaga, 1886, now L. trilineata (Bedriaga), of which Fitzinger's name iS now a jr. synonym). mildei, Amphisbaena, Peters, 1878, Monatsb. K. Preuss. Akad. Wiss. 1878: 778-781 (rejected in priority but not in homonymy contexts to protect Amphisbaena trachura Cope, 1885, now A. darwini trachura, of which Peters' name is now a jr. synonym). Minor, Testudo mydas, Suckow, 1798, Anfangsgr. Naturg. Thiere 3: S30) Meeqecteds an pritontty but not im homonymy “contiext’s to protect Thalassochelys (Colpochelys) kempii Garman, 1880, now Lepidochelys kempii (Garman), of which Suckow's name is now a jr. synonym). esses eprersirs, Allragator, Gray, VSl, “Syn. Rept.: 62 (an incorrect subsequent spelling of Crocodilus mississippiensis Daudin, 1801, now Alligator mississippiensis (Daudin), without nomenclatural status). MisciscapLtenois,. Crocodilus, Daudin, 1801, Hist. Nat. Rept. 2: ARIZ (an incorrect Or Lg ana d spelling, NioOn— es4hSit eve nomenclaturally, of Crocodilus mississippiensis Daudin, 1801, now Alligator mississippiensis (Daudin). fubrata, Rana). Houttuyn, 1787, “Anam. Mus... -Houtt..iIndex:, 19 (this work by Houttuyn has been suppressed for nomenclatural purposes (Opinion 380), hence the name is nomenclaturally non-existent and should not be assigned to any species; in addition the name is unidentifiable). molochina, Psammophis, Berthold, 1846, Mitt. Nachr. G.A. Univ. K. Geis’. / Wiss. Gottingen” Zool.Mus.\" Gottingen 1846: ARS Yes era (rejected in priority but not in homonymy contexts to protect Philodryas nattereri Steindachner, 1870, of which Berthold's name iS now a jr. synonym). monitor, acdemtva,, lannaeius,. 17587 Syst *=" Nat® 1S UE a el LO) eBne2x( OIL see eee in priority but not in homonymy aortas 1EG)) [SHAGIESCIE Stellio salvator Laurenti, 1768, now Varanus (V.) salvator (Laurenti), of which monitor is now a jr. synonym). Toleimeculata, Crotalus Juqubris, dan, 1863, Elenco. Sist..Ofidi: LA GeVecreds in sprlotiy but, MOL am Ihomonymy Contexts, tO 20 Protect: | Canidisiona jpiodiystwetagmiGope! jeleics,senowssGaionsquingis polystictus (Cope), of which Jan's name is now a jr. synonym). nasua, Vipera, Wagler, 1830, Nattirl. Syst. Amph.: 171 (rejected in priority but not in homonymy contexts to protect Heterodon dorbignyi Duméril, Bibron and Duméril, 1854, now Lystrophis dorbignyi (Duméril, Bibron and Duméril), of which nasua is now a jr. synonym) . neocwelsatnwenissis,, Proteus; Green pe sie, wd. Acad Nat Scie Philadelphia, 1:358 (rejected in priority but not in homonymy convextis vo protect) -Sallamandra jenigriimal iGreen, e257. now Ambystoma t. tigrinum (Green), of which Green's name, of 1818, is now a jr. synonym). nigeryyScyvtale; “Daudin, A803)" Hist “ NateimiGens Parties 93 Rept. -os42 (based on what is now known as Heterodon platirhinos Latreille, 1801, but rejected in both priority and homonymy contexts, to protect Scytale neuwiedii nigrum Duméril, Bibron and Duméril, 1854, now Pseudoboa nigra (Duméril, Bibron and Duméril); Daudin's name is now non-existent nomenclaturally). nigricollis, -Coluber,; Dwigubskij,.1832,. Opyt. Hstestv. »Istorid 13326 (rejected in priority but not in homonymy contexts to protect Coronella modesta Martin, 1838, now Eirenis modesta (Martin), of which Dwigubskij's name is now a jr. synonym). Oaxkacae, Kinostiennon,—slprltchand,: » LO 9 bneyicl. Twist lese 5 5y7/ (rejected in priority but not in homonymy contexts to protect Kinosternon oaxacae Berry and Iverson, 1980, of which Pritchard's name is now a jr. synonym). oce liad, Wacertay SHouttuyn sal Site Anime Muster HOUT tly slides: 24 (this work by Houttuyn has been suppressed for nomenclatural purposes (Opinion 380), hence the name is nomenclaturally non- existent and should not be assigned to any species; in addition the name is unidentifiable). oce}aitus,) Draco, Oken, 41816; Lehrb.wNatung 4433277 altthus work by Oken has been suppressed for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of any species). operculata, Siiren,. Beauvois, 17/99, Trans... Am. Phidios.. Soc. -4:277—- 281, figs. 1-4 a(mejected,, in ~praority. but» not ,anyihomonymy contexts, to, protect _jSialkamandray tivqaamiays Green, a lsiZo7= now Ambystoma t. tigrinum (Green), of which Beauvois' name is now a jr. synonym). Oryzicola, Berus, Oken, 1816, Lehrb. Naturg. 3:248 (this work by Oken has been suppressed for nomenclatural purposes (Opinion al 417), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of any species). eryzivorus, Coluber, Suckow, 1798, Anfang. Theor. Angew. Naturg. Thiere 3 Amphibien: 245) (rejected "in prLorbty sbutjonot in homonymy contexts to protect Boa reticulata Schneider, 1801, now Python reticulatus (Schneider), of which oryzivorus is now aajeeasSynonym)r. Op. 71463" oularsawa, Coluber, Bonnaterre, 1790, Tabl. Encycl. Meth. Trois Regnés Nature: 26 (rejected in priority but not in homonymy GCOnEeXES peeO, Protect, ‘Bod Pelt cullatay Schnicider,™ 1si0iyy) now Python reticulatus (Schneider), of which oularsawa is now a jr. synonym). Op. 1463. Papillosa, Rana, Houttuyn, 1787, Anim. Mus. Houtt. Index: 19 (this work by Houttuyn has been suppressed for nomenclatural purposes (Opinion 380), hence the name is nomenclaturally non- existent, and should not be assigned to any species; in addition the name is unidentifiable). pelamys, Hydrophis, Oken, 1816, Lehrb. Naturg. 3: 279 (this work by Oken has been suppressed for nomenclatural purposes (Opinion 417), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of any species). Detrefacktus, Crocodulus, Bererand, 795), DIGE., Unavi. HOSS. Propses Foss. Accid. 1: 183 (this work by Bertrand has been suppressed for nomenclatural purposes (Opinion 592), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of any species). nee! lphi PP PASCOMMOS Udo Ck mS 22). = AOC Ogata ssi: 179 (rejected in priority but not in homonymy contexts to protect Salamandra tigrina Green, 1825, now Ambystoma t. tigrinum (Green), of which Jarocki's name is now a jr. synonym). Planitia. TEestudo, Gmelin, 2789,.4n Linnaeus, sDop. Mus: ‘Natee Acad. Upsaliensis: 30 (rejected in priority but not in homonymy contexts to protect Amphisbaena cinerea Vandelli, 1797, now Blanus cinereus (Vandelli), of which Thunberg's name is now a jr. synonym). Salvaquardia, Stellio, Laurenti, 1768, Specimen Medicum: Si (rejected in priority but not in homonymy contexts to protect Tupinambis bengalensis Daudin, 1802, now Varanus bengalensis (Daudin), of which Laurenti's name is now a jr. synonym). sclerotica Elaphe, Smith, 1942, Copeia,) 1941: “135, 136 (Coluber subocularis Brown, 1901, now Bogertophis subocularis (Brown) ruled not invalidated by the sr. name Bascanion suboculare Cope, 1867, .a jr. synonym, of Masticophis m.. mentovarius (Duméril, Bibron and Duméril, 1854), hence Smith's name is a jr. objective synonym of Brown's name). semimembranacea, Testudo, Hermann, 1804, Observ. Zool. 219 (rejected in priority but not in homonymy contexts to Beoeaee Trionyx (Aspidonectes) sinensis Wiegmann, 1835, now Trionyx Sinensis Wiegmann, of which Hermann's name iS now a jr. synonym) . SUMPCHrast 1], -Hendcodnathus, Bocourt,, 1866, Missa sce e Mex. | (a0))7: 626-630), "pls 41, arg. 5. (rejected: ine pra omrty me bulies sO rs sn homonymy contexts to protect Ablabes chinensis Giinther, 1889, now Sibynophis chinensis (Giinther), of which Bocourt's name is now a jr. synonym). CErrestris, Testudo,~ Fermin,’ 1765,” Haste Nat. Hollande quanoxs. 51 (this work by Fermin has been suppressed for nomenclatural 28 purposes (Opinion) 660), hence the name does not exist nomenclaturally, and should not be cited in the synonymy of any species). tibiatrix, Hyla, Laurenti, 1768, Spec. Medicum: 34 (rejected in priority but not in homonymy contexts to protect Rana venulosa Laurenti, 1768, now Phrynohyas venulosa (Laurenti), of which H. tubiatreLxeus mMNoOw alla Synonym) timorensis, Python, Miiller, 1844, Verh. Natuurl. Gesch. Ned. Overz. Bez., Land- en Volkenk. (7): 211, 220 (segiecved an pErLOrlty but “Not “in homonymy contexts to protect Lzasis mackloti Duméril and Bibron, 1844, of which Muller's name is now a jr. synonym). timoriensis, Python, Miiller, 1857, Reizen en Onderzoekingen in den Indischen Archipel, gedaan op last der Nederlandsche Indische Regeering, tusschen de Jaren 1828 en 1836 2: 2 Grulked van incorrect subsequent spelling of Python timorensis Muller, 1844, hence without nomenclatural status). trimeresurus, Coluber dipsas, Oken, 1816, Lehrb. Naturg. 3:263 (this work by Oken has been suppressed for nomenclatural purposes (Opinion) 417), hence “the mame = does not exist nomenclaturally, and should not be cited in the synonymy of any species). oulC@cikcia, Chobadibtere, Uexeloelsl, Mishel, (Giles g Jeteicies 8 28 (Tschudi's usage, and all others joe Novia to the proposal of Eleutherodactylus unicolor Stejneger, 1904, are rejected in both priority and homonymy contexts, to preserve Stejneger's name, which would otherwise be a jr. homonym; Tschudi's name is actually referable to Eleutherodactylus and would, if not rejected, replace Leptodactylus inoptatus Barbour, 1914, now E. inoptatus (Barbour); Tschudi's name, as type of Cornufer, requires replacement in that role to leave the name Cornufer as long interpreted, through designation of Halophila vitiensis Girard, 1853, as type species, although that species is now generally referred to the genus Platymantis Giinther, 1858, along with all other species formerly referred to Cornufer; if Platyvmantis is split in the, future, Cornufer is available it needed; thus three names are protected by rejection of Tschudi's name). VENEENGCOSa; Emus, Gray,.1855;,.Gat. Shield, Rept. Coll Nomenel., UZ (ils) Qui eS)", ICZN. 1956. Opinion 417. Rejection for nomenclatorial purposes of volume 3 (Zoologie) of the work by Lorenz Oken entitled "Oken's Lehrbuch der Naturgeschichte" published in 1815-1816. Ope sDecl ys interns Comm. Zool.) wNomencili, slau) sla 2F ICZN. ILS) (ya Opanatonr = 519)2"- Rejection of Bertrand, 1763, Dictionnaire universel des fossiles propres et des fossiles accidentels. Bull. Zool. Nomencl., 18:114-120. ICZN. 1963. Opinion 660. Suppression under the plenary powers of seven specific names of turtles (Reptilia, Testudines). Bull. Zool. Nomencl., 20:187-190. 25 ICZN. 1966. Opinion 789. Rejection of the pamphlet by J. Hiibner, 1808, entitled Erste Zutrage zur Sammlung exotischer Schmetterlinge. Bull. Zool. Nomencl., 23:213-220. ICZN. 1974. Opinion 1024. Epirhexis Cope, 1866 (Amphibia: Salientia): Suppressed under the plenary powers. Budeise Zool sNomencl >, 6. (3) 130-132 MedvalbileyeRichardeVvasand od. D. D. Smith. 19875 Officials amalsies! tand Indexes of Names and Works in Zoology. International Trust for Zoological Nomenclature. London. Smith pe HObarteMen and ROzellal Be Smash. 1e)7/S) Synopsis of the Herpetofauna of Mexico. Vol. II. Lundberg; Augusta, West Virginia. Vamnz ollie aul Om Et 1977. An Annotated Bibliography of the Land and Fresh-water Reptiles of South America (1758-1975). Vol. reliTS 900) Mus. 4ool., Univ... Sac) Paulo; (Sao Paulo. CROSS-INDEXES The following cross-indexes will facilitate scanning the 200 names here treated for those pertinent to valid names. ies leatsic is divided into two groups: nomina clara (names whose allocation to known taxa iS apparent) and nomina dubia (names whose allocation to any given taxon is uncertain). NOMINA CLARA Indented names are to be found in the preceding account. CAUTION: not all indented names are synonyms of the valid names under which they appear; most are, but in some cases the valid name is simply discussed in the account for the invalid name. CAUDATA A. Family-Group Names Ambystomat idae Ambystomidae B. Genus-group Names Ambystoma Amblystoma Axolot Axolotus Philhydrus Phyllhydrus Siredon Sirenodon 26 Stegoporus Andrias Palaeotriton Proteocordylus Aneides Anaides Autodax Bolitoglossa Oedipus Palmatotriton Notophthalmus Diemichylus Diemyctelus Diemyctylus Notophthalma Notophthalmia Notopthalmus Siren Sirena Sirene C. Species-group Names Ambystoma tigrinum tigrinum neocaesariensis, Proteus operculata, Siren philadelphicus, Axolotus Aneides lugubris ereticauda, Triton Cryptobranchus alleganiensis alleghaniensis, Abranchus Gyrinophilus porphyriticus duryi lutescens, Triturus Plethodon cinereus erythronota, Salamandra Pleurodeles waltl funebris, Salamandra SALIENTIA A. Family-group Names Leiopelmatidae Leiopelmidae Liopelmatina Liopelmidae B. Genus-group Names Kassina Eremiophilus Leiopelma Liopelma Phrynohyas Qui Acrodytes Syrrhophus Epirhexis C. Species-group Names Eleutherodactylus inoptatus unicolor, Cornufer Eleutherodactylus unicolor unicolor, Cornufer Hyla boans maxima, Rana Phrynohyas venulosa Ebvataisa, HyAkai zonata, Hyla Platymantis vitiensis unicolor, Cornufer Rana grayi fasciata, Rana Rana perezi bosci, Rana GYMNOPHIONA A. Family-group Names Caeciliaidae Caeciliidae Cecilinia RHYNCHOCEPHALIA A. Family-group Names Sphenodontidae Hatteriidae Rhynchocephalidae B. Genus-group Names Sphenodon Hatteria Rhynchocephalus Sphaenodon TESTUDINES A. Genus-group Names Dermochelys Coriudo Dermatochelys Dermochelis Sphargis 28 Rhinoclemmys Chemelys B. Species-group Names Dermochelys coriacea mercurialis, Sphargis Emydoidea blandingii flava, Testudo meleagris, Testudo Kinosternon alamosae alamose, Kinosternon Kinosternon oaxacae oaxacae, Kinosternon Lepidochelys kempii minor, Testudo mydas viridi-squamosa, Testudo Macroclemys temminckii planitia, Testudo Rhinoclemmys punctularia dorsata, Testudo Trachemys ornata cataspila ventricosa, Emys Trionyx sinensis semimembranacea, Testudo Trionyx steindachneri californiana, Aspidonectes SAURIA A. Family-group Names Agamidae Agamoidea Stellionidae (2) Teiidae Tupinambidae B. Genus-group Names Agama Stellio Bachia Brachypus Chalarodon Tritropis Tropidogaster Draco Dracunculus Ichnotropis Thermophilus Mabuya Mabouia Mabouya 29 Mabuia Mabuya Phrynosoma Tapaia (2) Tapaja Tapaya (2) Tapayia Uromastyx Centrocercus C. Species-group Names Agama agama agama colonorum, Agama Agama bibronii bibronii, Trapelus (Psammorrhoa) Anguis fragilis colchica besseri, Anguis Bachia bicolor diglossis, Chirotes Chalarodon madagascariensis blainvillii, Tropidogaster Lacerta trilineata michahellesii, Podarcis Podarcis melisellensis fiumana punctato-fasciata, Lacerta muralis punctato-striata, Lacerta muralis Varanus bengalensis dracaena, Lacerta salvaquardia, Stellio Varanus (Varanus) salvator monitor, Lacerta AMPHISBAENIA A. Species-group Names Amphisbaena darwini trachura mildei, Amphisbaena Amphisbaena dubia dubia, Amphisbaena Amphisbaena fuliginosa amazonica dubia, Amphisbaena Blanus cinereus cinereus reticulata, Amphisbaena SERPENTES A. Genus-group Names Ahaetulla Dryinus Dryophis Passerita 30 Anilius Ore tae is Bases Cobra Boa Constrictor Boiga Ibiba Cemophora Amplycephalus Clonophis Cora Dasypeltis Anodon Dendrelaphis Tachyophis Dromophis Philodendros Philodendrus Enhydris Enhydrus Leptophis Ahaetulla Dendrophis Leptotyphlops Typhlina Lystrophis Rhinostoma Ophidioniscus Ophidion Ophiophagus Dendraspis Hamadryas Pareas Amplycephalus Phimophis Rhinosimus Sibon Asthenognathus Simophis Rhinaspis Spalerosophis Sphalerosophis B. Species-group Names Boa constrictor formosissimus, Constrictor Bogertophis subocularis sclerotica, Elaphe Cemophora coccinea doliatus, Coluber CrotalusVatrox Ccinereous, Crotalus Sil Crotalus horridus cincolor, Crotalus durissus concolor, Crotalus durissus Crotalus polystictus multimaculata, Crotalus lugubris Crotalus viridis decolor concolor, Crotalus durissus Drymarchon corais melanurus melanocercus, Drymarchon corais Drymobius margaritiferus chiametla, Coluber Drymobius margaritiferus fistulosus chiametla, Coluber Eirenis modesta nNigzercollsts,, Coluber Elaphe scalaris elaphis, Coluber quater-radiatus, Coluber Hemorrhois viridiflava carbonaria melanepis, Coluber Heterodon platirhinos niger, Scytale Liasis mackloti timorensis, Python timoriensis, Python Lystrophis dorbignyi nasua, Vipera Macropophis barbouri vertebralis, Leptophis Ninia atrata atratus, Coluber Philodryas nattereri molochina, Psammophis Philothamnus irregularis caesius, Coluber Psammophis subtaeniata bilineatus, Psammophis moniliger Pseudoboa nigra niger, Scytale Python reticulatus oryzivorus, Coluber oularsawa, Coluber Sibynophis chinensis sumichrasti, Henicognathus Simophis rhinostoma proboscidea, Rhinostoma proboscidea, Rhinostoma (Rhinaspis) Storeria occipitomaculata leberis, Coluber Vipera ursinii renardi foetidus, Coluber 32 CROCODYLIA A. Species-group Names Alligator mississippiensis alligator, Lacerta lucius, Crocodilus (Alligator) mississipiensis, Alligator mississipiensis, Crocodilus Crocodylus palustris indicus, Crocodilus vulgaris NOMINA DUBIA A few of the invalid names listed in the preceding section are nomenclaturally non-existent, but are included there because they have commonly been included in synonymies. Thirty-seven other names, not commonly cited in synonymies, and non-existent nomenclaturally (hence not obliged to be cited), follow, with author and date. SALIENTIA Bufo cruciger Oken, 1816 Rana coerulea Houttuyn, 1787 Rana mitrata Houttuyn, 1787 Rana papillosa Houttuyn, 1787 Rana verrucosa Houttuyn, 1787 TESTUDINES Testudo terrestris Fermin, 1765 SAURIA Discosomus Oken, 1816 Petrodactylus Oken, 1816 Pterodactylus Oken, 1816 Seine? Oken, 18116 Scincorum Oken, 1816 Zygnis Oken, 1816 Anguis granulatus Houttuyn, 1787 Bipes lepidopus Oken, 1816 Draco ocellatus Oken, 1816 Lacerta areolata Houttuyn, 1787 Lacerta fasciata Houttuyn, 1787 Lacerta grisea Oken, 1816 88 Lacerta ocellata Oken, 1816 Scincus galliwasp Oken, 1816 Stellio graecus Oken, 1816 stellio italicus Oken, 1816 AMPHISBAENIA Propus Oken, 1816 Amphisbaena flavescens Houttuyn, 1787 Amphisbaena marmorata Houttuyn, 1787 SERPENTES Berus Oken, 1816 Draco Oken, 1816 Herpeton Oken, 1816 Berus oryzicola Oken, 1816 Coluber cupreus Houttuyn, 1787 Coluber dipsas trimeresurus Oken, 1816 Coluber jacapara Houttuyn, 1787 Crotalus durissus concolor Jan, 1859 Hydrophis pelamys Oken, 1816 Trigonocephalus lancifer Oken, 1816 CROCODYLIA Crocodilus Bertrand, 1763 Crocodilus petrefactus Bertrand, 1763 ssagmn eigaiss ipplenwae’ or, Lacerta we peluetris ettndtcur, Créeodi tue vulgar a pigtural Ly not-existent, ey’ ve gonmamaly been joicluded «i newed, fo commonly. cited (hn Bye Momengleturaliy (hence pet obliged JERS natnor 265 date arat : aa oe Ce == , 7 re “HoAtteyn, +18? hevttcuyr, 1767 _ ieee. 7 Heuttuyn, 1737 — €80F yhosx3298 tavt sas: r " : TENTUDINES 7 ie Sesh ids \Celecercis Fermin, 2765 7 i: 3 “a | SAUPIA ae Discasupks Okan) 2446 - | \¢ ; Fetrodscivils Olen, 1935 eee Mean, (81t Anguis @sanuloius Nouvtuyn, 17387 Rapes Jéepiecgur Chan, 1816 Disco Oran, 1616 latishh Gsenieea fevteuyn, 176) featiana Movttuyns 1787 : giland Okeny 38L6 A CATEGORIZATION AND BIBLIOGRAPHIC SURVEY OF PARENTAL BEHAVIOR IN LEPIDOSAURIAN REPTILES WS) wr Uw SWANN . AN pvt ANY Mm <7 AS 0 NAN \\\ \, Wy ae \ WA \\ YP. LOUIS A. SOMMA Department of Zoology University of Florida SMITHSONIAN HERPETOLOGICAL INFORMATION SERVICE No. 8l 1990 SMITHSONIAN HERPETOLOGICAL INFORMATION SERVICE The SHIS series publishes and distributes translations, bibliographies, indices, and similar items judged useful to individuals interested in the biology of amphibians and reptiles, but unlikely to be published in the normal technical journals. Single copies are distributed free to interested individuals. Libraries, herpetological associations, and research laboratories are invited to exchange their publications with the Division of Amphibians and Reptiles. We wish to encourage individuals to share their bibliographies, translations, etc. with other herpetologists through the SHIS series. If you have such items please contact George Zug for instructions on preparation and submission. Contributors receive 50 free copies. Please address all requests for copies and inquiries to George Zug, Division of Amphibians and Reptiles, National Museum of Natural History, Smithsonian Institution, Washington DC 20560 USA. Please include a self-addressed mailing label with requests. INTRODUCTION Parental behavior is a significant aspect of the life history of a wide variety of animal taxa. Parental behavior is common to the vertebrate classes Osteichthyes (Blumer, 1979, 1982; Perrone and Zaret, 1979; Baylis, 1981; Gittleman, 1981; Gross and Sargent, 1985), Amphibia (McDiarmid, 1978; Wells, 1981; Nussbaum, 1985; Duellman and Trueb, 1986), and universal within the Aves (Kendeigh, 1952; Skutch, 1957, 1976; Welty, 1982; Silver et al., 1985) and Mammalia (Kleiman and Malcolm, 1981; Dewsbury, 1985; Kleiman, 1985; Malcolm, 1985). In contrast, relatively few reptiles exhibit parental behavior (Tinkle and Gibbons, 1977; Shine and Bull, 1979; Shine, 1985, 1988). Parental behavior is common in living crocodilians (Greer, 1970, 1971; Lang, 1987; Shine, 1988) and also may have been common in extinct archosaurs (Horner and Makela, 1979; Coombs, 1982; Horner, 1982, 1984, 1987; Horner and Gorman, 1988) and cynodonts (Graves and Duvall, 1983; Duvall, 1986). Accounts of parental behavior in four turtle species (Gopherus agassizii: Barrett and Humphrey, 1986; G. flavomarginatus: Carr, 1952; Janulaw and Appleton cited in Morafka, 1982; Appleton, 1986; Ernst and Barbour, 1989; Manouria emys: Louwman, 1982; McKeown et al., 1982; Trachemys stejnegeri malonei: Hodsdon and Pearson, 1943) are remarkable, because turtles generally lack any form of parental behavior (Shine, 1988; Ernst and Barbour, 1989). The purpose of this review is to survey the various categories of parental behavior reported for lepidosaurians (lizards, snakes, amphisbaenians and a rhychocephalian) and to provide an extensive bibliography as a guide to current and future researchers. The phrase "parental care" refers to all nongametic and postfertilization contributions of parents to the survival of their offsprings (Wittenburger, 1981; Blumer, 1982) and is construed by some (Williams, 1966; Baylis, 1981; Keenleyside, 1981; Gross and Sargent, 1985; Congdon, 1989; Spotila and O'Connor, 1989) to include viviparity and other physiological contributions. I use the phrase "parental behavior" to limit the scope of this survey to all behavioral contributions by the parent to offspring survival after oviposition or parturition. Behaviors associated exclusively with oviposition and nest construction are not included in this survey; they are probably common to most oviparous lepidosaurians (Hahn, 1909; Hilzheimer, 1910; Blanchard, 1933; Carl, 1944; Carpenter, 1966; Platt, 1969; Rand and Rand, 1976; Duvall et al., 1979; Jones and Guillette, 1982; Green and Pauley, 1987). The term "brooding" describes behaviors of the parent while attending the nest and progeny (sensu Somma, 1988; also see Pope, 1961; Peters, 1964; Carpenter and Ferguson, 1977). Thus, brooding does not include territorial, nest-site defense wherein the parent remains at a distance from the nest, and not in or on the nest or progeny, as seen in some iguanine and gekkonid lizards, and the tuatara. HISTORICAL ASPECTS The oldest conceivable documentation of parental behavior ina lepidosaurian is in the book of Isaiah, 34:15 (McDowell et al., 1982) written some time between 745-350 BC (Asimov, 1968). In this account, the Hebrew word "lilith," at one time interpreted as an owl (Strong, 1961; Asimov, 1968), is referred to as the "arrow snake." This vernacular name may have referred to the boid Eryx jaculus (Topsell, 1608; White, 1954). Since E. jaculus is viviparous, the snake reputed to "lay eggs and hatch, and gather them under her shadow" (McDowell et al., 1982) may refer to an oviparous snake or be erroneous, if not simply a fable. Much of the pre-Twentieth century natural history literature indicates that many authors believed parental behavior was universal in snakes (Aristotle, [d. 322 BC]; Nicander of Colophon [135-133? BC]; Gesneri, 1551- 1587; Topsell, 1608; Chateaubriand, 1827; Sundowner, 1895, 1902). The Second Voyage of Sinbad the Seaman, written in the 8th century AD (Burton, 1885-1888), is an example of early fiction that mentions a giant snake (undoubtedly a python) brooding its eggs. The "cockatrice" or "basilisk" was reputed to brood her eggs (Gesneri, 1551-1587; Topsell, 1608). It is likely that this mythical beast was a fantastical description of a cobra, either Ophiophagus hannah or a species of Naja (White, 1954; also see descriptions by Pliny the Elder, [d. AD 79]; Gesneri, 1551-1587; Topsell, 1608). Similarly, brooding was attributed to the "asp" (= Naja haje?) and the "dipsas" (= Bungarus sp.?; White, 1954) by Nicander. Modern documentation confirms that these taxa brood their eggs (Table VI). Snakes have long been credited with the ability to swallow their young to protect them from danger (Topsell, 1608; Carver, 1778; Mease, 1807; Holmes, 1823; Hunter, 1824; Chateaubriand, 1827; Gosse, 1851; Rivers, 1874; Stanley, {19??]; Burroughs, 1908; Meek, 1946). Earliest accounts of this behavior are found in hieroglyphics attributed to the ancient Egyptians circa 2300 BC (Speck, 1923). So prevalent was (and still is!) this belief, that it was incorporated into early fiction, including the pre-Elizabethian poem, The Faerie Queene (Spenser, 1590). This alleged behavior, attributed not only to snakes but also to the lizard, Lacerta vivipara (Hopley, 1882), has been reviewed and discussed by numerous authors for more than 300 years (Browne, 1646; White, 1787; Hopley, 1882; Noble, 1921; Speck, 1921, 1923; Schmidt, 1929; Ditmars and Bridges, 1937; Angel, 1950; Klauber, 1972; Russell, 1983; Shine, 1988). Despite many inquiries into the plausibility of this behavior, no scientific evidence exists for its occurrence (Klauber, 1972; Shine, 1988). Among saurians, Scincus scincus was reputed to care for its eggs and young (Gesneri, 1551-1587; Topsell, 1608), but no modern observations support this contention (Table V). Hoy (1883) suggested that all lizards brooded their eggs. In all likelihood, his statement is based upon observations of Ophisaurus attenuatus and Eumeces septentrionalis; these are the only egg-brooding lizards that he actually observed (Hoy, 1883). The suggestion that an amphisbaenian broods its eggs (Gesneri, 1551-1587; Topsell, 1608; Aldrovandi, 1640; reviewed in Druce, 1910), has not been verified. CATEGORIES OF PARENTAL BEHAVIOR The various categories of parental behavior, as reported in the literature, are listed below. The symbol in parentheses identifies the categories used in Tables I and II. The literature sources are provided in Tables V and VI. Coil around brood (C): The attendant parent remains coiled around or covers the brood with its body, presumably creating a physical buffer or barrier between progeny and the external environment. This is the most common form of parental behavior. Nest constructed and maintained (NC): A burrow or brood chamber is constructed by the parent and maintained while attending progeny. Although this usually involves digging a depression or burrow in the substrate, Ophiophagus hannah is known to maintain a relatively complex nest chamber constructed from surrounding vegetation (Wasey, 1892; Oliver,1956; Leakey, 1969; Whitaker, 1977). Defense of brood (D): Progeny are aggressively defended by parent in the presence of conspecifics or heterospecifics. Passive protection (PP): Neonatal vipers may accrue protection from the venomous female, without her exhibiting any overt signs of aggressive behavior (i.e., Crotalus horridus: W. Martin, pers. comm.). Thermoregulation (T): Attendant parent uses its body to maintain a relatively constant incubation temperature for developing eggs. Most, perhaps all, pythonines are able to become low-grade endotherms while brooding through "shivering thermogenesis," thereby raising the temperature of the female's body and developing eggs above that of ambient conditions (Vinegar et al., 1970; Harlow and Grigg, 1984; Shine, 1988). However, it has been suggested that shivering thermogenesis is not practiced by all species of pythons (Vinegar et al., 1970; Ellis and Chappell, 1987; but see [Orlov], 1986; Shine, 1988). It is also possible that an attendant parent could (1) provide a passive thermal barrier between eggs and the external environment with its body or (2) bask in the sun and transfer radiantly absorbed heat from its body to its eggs (Medsger, 1919, 1932; Noble and Mason, 1933; Cogger and Holmes, 1960). Python eggs, of at least two species, that are not brooded, and subsequently incubated at lower temperatures, take longer to hatch and exhibit a higher rate of developmental anomalies (Vinegar, 1973, 1974; Branch and Patterson, 1975). Hydroregulation (H): Even though is has never been demonstrated experimentally, hydroregulation has been inferred from some squamates (Fitch, 1954; Somma, 1985b; Bels and Van den Sande, 1986; [Orlov], 1986; York and Burghardt, 1988; Somma, 1989b; Somma and Fawcett, 1989). In addition, tenuous evidence suggests that two species of snakes wet their bodies with water and then lay over the eggs, thereby reducing desiccation (Elaphe obsoleta: J. Lombard, pers. comm.; Trimeresurus wiroti: Mehrtens, 1987). False brooding (FB): Many species of pythons continue to brood when their clutches are removed prior to hatching (T. Miller, pers. comm.), and a Python molurus has brooded without ovipositing (J. S. Foster, pers. comm.). The nongravid female P. m. bivittatus housed with two gravid conspecifics brooded when the gravid females oviposited and brooded their eggs. This "false brooding" consisted of the python laying in a conical coil and exhibiting shivering thermogenesis (J. S. Foster, pers. comm.; Somma, pers. observ.). Oophagy (00): The parent eats eggs or aborted ova. This is a form of parental behavior because it may prevent microbial infection of viable, healthy eggs from adjacent infected eggs (Groves, 1982; Somma, 1989a) or prevent detection of progeny by predators using olfaction (i.e., detection of chemical cues released from rotting eggs or aborted ova) to locate food (Tinkle and Gibbons, 1977; Groves, 1982; Shine, 1988). Earliest observations of this behavior may be found in Hindu scriptures dating back to 600 BC (Rao, 1957). Removal of nonviable eggs from nest (RE): Nonviable eggs are removed from the nest, presumably at a safe distance from viable eggs. This behavior has been reported only in Python molurus (Griehl, 1982) and perhaps occurs in Gerrhonotus liocephalus (Greene cited in Tinkle and Gibbons, 1977). Parental care of neonates (CN): A parent remains with and expresses parental behavior toward neonates after hatching or parturition. Neonates assisted during hatching or parturition (AN): Attending parent assists neonates from eggs, extraembryonic membranes or nest chamber. In Eumeces obsoletus and E. septentrionalis, the mother licks amniotic fluid from neonates' bodies after hatching (Evans, 1959; Somma, 1987c). Manipulation or retrieval of eggs (ME): Eggs are manipulated within the nest or moved to a new nest site. Eggs that have been removed from the nest may also be retrieved. Communal care of eggs (CC): Eggs may be deposited in a communal clutch and attended by several unrelated parents. In some situations, this behavior could be interpreted as alloparental care (Banks and Schwaner, 1984; Somma, 1987a). Neonatal feeding facilitated (FN): A brooding female E. obsoletus avoided taking food items in deference to its young, waiting for them to finish before she fed (Evans, 1959). This behavior has not been observed in other reptiles. Normally does not exhibit parental behavior (NPB): Parental behavior has been reported (sometimes reliably) for individuals of species that normally abandon their progeny. Details unknown (DU): Details of parental behavior are not known or not reported. Reliability uncertain (RU): Reliability of report regarded as uncertain or possibly unreliable due to paucity of information provided or observed by the author. Erroneous documentation (E): Report considered unsubstantiated due to inadequate information for a species normally lacking parental behavior. DISCUSSION Over 6140 extant species of lepidosaurian reptiles are currently recognized (Bellairs, 1986); of these, parental behavior has been reported for 210 species. Only 148 species (82 lizards, 65 snakes, and Sphenodon) are represented by reliable documentation or approximately 2.4% of all lepidosaurians (Table III). These species represent 17 families, excluding the unsubstantiated documentation for the Hydrophiidae, Typhlopidae and the unidentified amphisbaenid family (Tables I, II, III). Parental behavior is common in Eumeces, Phelsuma, Uromastix, anguids, xantusiids, iguanines, oviparous boids, southeast-Asian elapids, oviparous viperids, Elaphe, Farancia, and Psammophylax (Tables I and II). In most species (96%) parental behavior is entirely maternal; however, paternal and biparental behavior have been confirmed (Table IV). The literature in this survey (Tables V and VI) demonstrates a paucity of experimentally obtained data on parental behavior in lepidosaurians (but see Noble and Mason, 1933; Hutchison et al., 1966; Vinegar et al., 1970; Van Mierop and Barnard, 1976a, 1978; Hasegawa, 1985, Somma, 1985b; [Orlov], 1986; Ellis and Chappell, 1987; York and Burghardt, 1988; Graves, 1989; Somma and Fawcett, 1989; Vitt and Cooper, 1989; Guillette et al., in review). The majority of the literature is descriptive or anecdotal, and the adaptive functions of this life history trait remain largely conjectural. Furthermore, the only ecological/evolutionary analysis devoted solely to parental behavior in reptiles is provided by Shine (1988; see Lillywhite, 1988). ACKNOWLEDGEMENTS Obtaining the literature for this survey would have been impossible without the superlative library skills of J. Mundell, L. Valentine and the late M. Covault (Interlibrary Loan Department, University Library, University of Nebraska at Omaha), D. Beaubien, B. Gilbert and K. Harmon (Interlibrary Loan Department, Marston Science Library, University of Florida), and S. Tergas (Interlibrary Loan Department, Main Library West, University of Florida). I am also grateful those who provided or informed me of important references; they are E. J. Bredin, R. L. Burke, D. B. Carter, B. Clark, P. A. Cochran, H. Ehmann, D. Ferraro, L. A. Fitzgerald, J. S. Foster, H. W. Greene, L. J. Guillette, Jr., A. Hagedorn, T. Heaton-Jones, K. Horikoshi, T. R. Johnson, G. Kattan, J. W. Lang, J. F. Lokke, S. A. Minton, Jr., L. D. Moehn, J. C. Murphy, L. Nico, J. G. Robinson, H. M. Smith, N. M. Somma, S. F. Somma, S. Stewart and S. J. Walsh. I wish to thank all of the individuals who allowed me to cite their unpublished observations as personal communications. I am indebted to D. Haney, A. Hensley and K. Horikoshi for translating the French, German and Japanese literature, respectively. I sincerely thank H. B. Lillywhite for his insightful comments and criticism of the manuscript. Special thanks are extended to R. Shine for his helpful comments and opinions. I owe my sincerest gratitude to J. D. Fawcett for his assistance, use of his extensive herpetological library, comments and proofreading of the manuscript, his boundless encouragement and enthusiasm for this project. Lastly, I thank J. Matter for allowing me to use his word processor and G. Kiltie for her skillful preparation of this manuscript. The cover illustration depicts a female Eumeces s. septentrionalis from Nebraska brooding eggs. I thank John F. Lokke for creating the original artwork from which the illustration was copied. TABLE I Distribution of parental behavior in lizards, amphisbaenians and a rhynchocephalian Oviparous Maternal, Type of or paternal or Taxa parental behavior viviparous biparental Agamidae Leiolepis belliana RU, CN O B? Phrynocephalus sp. RU, CN (@) B Uromastix aegyptius D (@) M U. ornatus D, CN 0) M Anguidae Barisia imbricata AN V M Diploglossus bilobatus C 0 M D. delasagra C O P Elgaria coerulea AN V M E. multicarinata GLENG DAHeC 0) M Gerrhonotus liocephalus C5; 2007, SRE? (0) M Mesaspis moreleti AN V M Ophisaurus apodus Card 6) M O. attenuatus C, 00 0) M O. compressus ei(C 0 M O. gracilis C 0 M QO. harti C ) M QO. ventralis C, ME, 0O O M Cordylidae Cordylus cataphractus CN V B C. giganteus CN V M Gekkonidae Ailuronyx seychellensis CDE LOO; JAN? 0 M Chondrodactylus angulifer RU, D? 0 M Eublepharis macularis RU, D, NPB (0) M Gekko gecko CD) -NPB?, 0 M, @B G. petricolus Gi, D200 (0) MireP G. smithii RU, DU 0 M Hemidactylus turcicus Hemiphyllodactylus typus Naultinus grayi Phelsuma borbonica dubia flavigularis lineata madagascariensis P. standingi Phyllodactylus lanei Ptychozoon lionotum Teratoscincus scincus lolrulrulro Iguanidae Amblyrhynchus cristatus Brachylophus fasciatus B. vitiensis Conolophus pallidus C. subcristatus Crotaphytus collaris Cyclura carinata C. cornuta C. cychlura (Ge seibloyiale Iguana iguana Phrynosoma douglassi Sauromalus varius Sceloporus undulatus Q Lacertidae woh oR o io) RU, NPB 2) = See elec ie] OQ 2 Acanthodactylus scutellatus RU, CN Lacerta viridis Scincidae Calyptotis scutirostrum Corucia zebrata Cyclodina pseudornata Egernia cunninghami E. striata E. whitii Emoia cyanura Eumeces anthracinus callicephalus chinensis copei egregius elegans . fasciatus . inexpectatus . laticeps latiscutatus lynxe multivirgatus obsoletus eaiiesiiesiiesi ies ies ileal ies seal ies ies] ie3) @, De NGs CC (s EN. 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