oo 3 9 LO te ons eer we thn Oa, Be Abo SAR tk de a OF 0 na : “ re : Koders Ate fre eome, tees “> SNe SOT eRe Neo . ; Pte wor ke ve MR om4 onearoy tet dsien oe Se See eee eg Pht emmacer oe Se arene Ate a Ne Metin Ie Nocare WS Nemes $5 SNS Setar Se To UNIVERSITY QF ILLINOIS LIBRARY AT YRBANA-CHAMPAIGN BIOLOGY 4 F , > 5) VYUL A tt Pag Oe ‘ ' ze ms * i i i * ¢ ' 7 ‘ wl , 7 = ed 7 i . Ee ; a > KE = nae Li a i is 7 a ; . i : : ; I ‘ Com ; = 4 ‘a p 4 ; cf ‘ , ¥ £ - 2) ee 7 ae 4 x % + = 4 * + - »s i 7 co 7 ae 7 Bs G ; “ : ‘ a " 7 7 ei : a anew, A oes ; 7 U Ox; : ') Bin Ces et ; a ‘ i are ei ; y vi f 1h ee ee vee Y hae Oe 5 ae aN ; i 2 a ptt FIELDIANA Zoology Published by Field Museum of Natural History Volume 72, No. 2 December 16, 1977 Amphisbaena medemi, An Interesting New Species from Colombia (Amphisbaenia, Reptilia), with a Key to the Amphisbaenians of the Americas CARL GANS DIVISION OF BIOLOGICAL SCIENCES UNIVERSITY OF MICHIGAN, ANN ARBOR AND SANDRA MATHERS DIVISION OF BIOLOGICAL SCIENCES UNIVERSITY OF MICHIGAN, ANN ARBOR SUMMARY Amphisbaena medemi, new species, is a small amphisbaenian from extreme north- ern South America. The species belongs to the grouping of small, four-pored animals and is suggestive of a stage intermediate between species from the mainland of South America and from the Antilles. The paper includes a key to all recognized forms of American amphisbaenians. The amphisbaenians of northern South America (cf. Gans, 1967) fall into two obvious groups. The first includes two large species, Amphisbaena alba and Amphisbaena fuliginosa, wide-ranging forms that occur across the Amazon basin and far to the south (cf. Gans, 1962a; Vanzolini, 1951). The second includes a series of five much smaller species, all with relatively restricted ranges, perhaps parapatric but certainly included within the ranges of the larger forms. The character states of these small forms suggest some di- versity; however, they all share such characteristics as fewer than 18 dorsal and fewer than 20 ventral segments to a mid-body annu- lus, a bluntly-rounded head with relatively large and regular shields and body annuli between 205 and 250. Four of the five species show four precloacal pores. If the poorly characterized Amphisbaena US ISSN 0015-0754 wa OGY LIBRARY \ The Library of the OLE URRILE HAE : Publication 1271 101 © 21 JAN 27 1978 . te Indis ES \ yuiversity of eee ee, 10, at (Irhana-Cean 22 FIELDIANA: ZOOLOGY, VOLUME 72 stejnegeri (known from two damaged specimens) is eliminated, the clustering of character states becomes even more marked. The assemblage is perhaps most interesting because one of its members, Amphisbaena gracilis Stauch, has been shown to repre- sent those conditions among all mainland forms that appear closest to the character states of the nine species of Amphisbaena inhabit- ing the Greater Antilles and adjacent islands (Sponga and Gans, 1971). Consequently, it is of some interest here to report on the dis- covery of yet another small mainland species that not only fits into the continental grouping but shows further affinity to the Antillean populations. We are grateful to Mr. Hymen Marx and Dr. Harold Voris, Field Museum of Natural History (FMNH), for making these specimens available for study and take pleasure in naming this new form after Dr. Fred Medem, collector of most of the specimens, in recognition of his substantial contribution to our knowledge of the herpetofauna of Colombia. Supported by National Science Foundation grant BMS 71 01380. Amphisbaena medemi new species Holotype.—FMNH 165245, taken between 1 and 10 December 1965 at Cienaga de Amajehuevo, Canal, Depto. Atlantico, Colombia by Federico Medem H. Paratypes.—FMNH 165243-165244, 165246-165247, taken be- tween 1 and 10 December 1964 with the holotype, FMNH 165248- 165250, taken on 15 January 1965 by C. A. Velasquez, and FMNH 165251-165257 taken on 31 January 1965 by F. Medem; all from the same locality. Diagnosis. — A small to medium-sized form of Amphisbaena with the nasals totally separated or reduced to medial point contact by the ascending rostral which generally contacts the frontals. Speci- mens often have azygous fusion of postocular and parietal head scales; three equally sized supralabials; one medium, one large, and one small infralabial on each side; two postgenial and no postmalar rows of chin shields. There are 230-235 body annuli; 17-18 caudal annuli with the autotomy constriction falling on the fifth to seventh postcloacal annulus; 16 dorsal and 18 ventral segments per midbody annulus; and four large circular precloacal pores in both sexes. The specimens are brown dorsally, fading to a lighter color ventrally. The color is densest in the rectangular centers of each segment, with the dark area decreasing along the sides and dropping out by seg- WY, “pessnosip ule1ey suvtusegsiydure [[eus Jay30 ayy Jo Sesues ay} 0} UOIZLAI Ul MWapam Duangsiyduy Jo Aye] BY MOYS 0} deut Jo yoJeHG “T “OLY HU1JOZUBA “YD e abaulajs @/3Z0/ Yj s1j19e 16 y 1/ja4inds vr /Wapau 23 Fic. 2. Amphisbaena medemi. Dorsal (top), lateral (middle), and ventral (bottom) views of FMNH 165245 show segment arrangement. The line equals 1 mm. to scale. (Don Luce, del.). 24 Fic. 3. Amphisbaena medemi. Dorsal, lateral, and ventral views of FMNH 165245 to show pigmentation, and the sculpturing of the surface. 25 Fic. 4. Amphisbaena medemi. Details to show variation. Frontal views of FMNH 165253 (a), and 165249 (b) to show extremes in the kinds of contact between rostral tip and frontals. Views of the nuchal region of FMNH 165244 (c) showing fusion of postocular to parietal on the right side and of FMNH 165256 (d) showing bilateral fusion. (Don Luce, del.). The line equals 1mm. to scale. 26 GANS & MATHER: AMPHISBAENIANS 27 Fic. 5. Amphisbaena medemi. Dorsal view (left), lateral view (middle), and ven- tral view (right) of FMNH 165243 at mid-body to show proportions and pigmen- tations of segments. ments in the neck and caudal regions. Many specimens show a light nuchal band. Description. — Figures 2 through 7 show aspects of segmentation, pigmentation and body proportions. Figure 8 presents a scatter dia- gram of tail lengths versus snout-vent lengths. The raw data for the type series are given in Table 1. This is a small to medium-sized species of Amphisbaena, showing a brownish dorsal and a lighter ventral coloration. The tip of the snout and the nuchal region tend to be lightened, while the dorsal, lateral, and posterior surfaces of the tail are densely pigmented. The lateral limits of pigmentation extend onto the ventral surface of the tail so that only the medial two to four rows of segments are light- ened. The intersegmental and interannular sutures are always lighter than the scale centers. Rostral, nasals, and superlabials as well as the chin shields tend to be unpigmented. Two or three annuli between the second and seventh may be faded or may lack pigmen- tation producing a light band across the nuchal region. The dorsal surface is more or less uniformly pigmented with only slight emphasis of the segmental centers. The two middorsal rows of segments are significantly darkened. One or two rows below the 0 NSS oO tao | ~~ (ai JoyueIq [ley 8ST FI 8 v &/€ € j 81/91 LT (S) v 9ST : 8 v &/€ € j 8I/9T 5 s : PST 8 8 v &/€ v j 81/91 LT (9) v : = €/€ v S 8I-9T/9T-ST : : - SST vI L v &/€ v j 8I-LT/91-ST 8I (9) € O9T LT 8 v &/€ v j 81/9T-ST 8I (9) € 8ST LI 9 v €/€ € td 8I/9T-ST LI (9) v 6ST vI L v &/€ v j 8I/91-ST 8I (L) € O9T LT L v &/€ g j 81/91 LT (9) g 8ST ST L v &/€ g (6 8I/91-F1 LT (9) v 6IT v1 8 v &/€ g € 8I/9I-ST LT (9) v OLT al L v &/€ g j 8I-LI/9T 8I (L) g T9T ; 9 v &/€ g j 81/9T = brl ST 9 v &/€ v j 81/91 LT (9) g qUaeA speoRo[D s[eovopeig sei0og elqeieijuy [etues [etues ye17u9 A, jepneg Aulojoyny [esszeT -ynouS =—s -980d /terqepeidng = -4s0q pugZ_—-480q 1ST /{es10q ‘WU - SJUSUBAINSBO |] ‘MapaU “V Jo suaulIdeds a] qeIIBAR ay} 1OJ SoqBYS JOJIVIVYO "I ATAVL 4 Apog LGZS9T 9S¢99T SSCG9T VSCS9T €SCS9T 6S6S9T OSZS9T 6PZS9T 8PZS9T L¥GS9T 9VZS9OT GPSS9T vVPSCSOT €PZS9T 28 GANS & MATHER: AMPHISBAENIANS 29 lateral sulcus the pigmentation becomes restricted to the segmental centers and these may become very small and faint by the fifth or sixth ventral row from each side. The three or four annuli anterior to the cloaca and the midventral segments of the caudal annuli show checkerboarding with the pigmentation appearing to drop sharply and often asymmetrically on individual segments. The head segmentation is characterized by a caudad projection of the dorsal tip of the rostral which partially or totally separates the nasals from medial contact and in some specimens inserts briefly between the prefrontals, and by the generally azygous fusion of parietal and postocular head scales; yet lacks other major fusions. The head is dorsoventrally compressed; the slightly elongated snout is dorsally arched but is of a horizontally oval cross-section. The rostral tip extends beyond the lower jaw and the temporal bulges noticeably posterior to the eye. There is some indication of nuchal constriction and slight reduction of the diameter of the nu- chal region. The trunk is approximately the same diameter along its length and is compressed dorso-ventrally representing at midbody a horizontally oval cross-section. The rostral is visible from above, slightly larger than the first su- pralabial, and of the same area as the nasals. Pairs of separated nasals, large prefrontals, medium-sized frontals, and smaller, quad- rangular to pentagonal parietals form a sequence along the dorsal surface of the head. The posterior tips of the frontals insert along the anterior suture between the parietals. Only four specimens have discrete parietals, in six that of the right side is fused to the en- larged postocular, in one fusion is on the left side and two show both sides fused. The diamond-shaped oculars are in anterior contact with the third supralabials and the large postocular. Each of three subequal supralabials is slightly larger than the ocular. The post- supralabial (just posterior to the angulus oris) abuts dorsally on the very large first temporal (which may be in point contact with the ocular). The postsupralabials, temporals, postoculars, and frontals apparently comprise the dorsal segments of the first body annulus. The four middorsal segments of the second body annulus are irregu- larly enlarged and lie posterior to the parietals, which thus seem to represent a medially intercalated pair of segments. The eye is clear- ly apparent. The mental is slightly narrower than the rostral, its area equiva- lent to that of the first infralabials which are followed by large sec- O10 LOJo Fic. 6. Amphisbaena medemi. Ventral view of cloaca and tail of FMNH 165257 to show segmental proportions. The line equals 1 mm. to scale. (Don Luce, del.). 30 FIG. 7. Amphisbaena medemi. Dorsal and ventral views of the tail of FMNH 165243 to show pigmentation and overall proportions. 31 32 FIELDIANA: ZOOLOGY, VOLUME 72 ond infralabials and smaller rectangular third infralabials. The ovate, posteriorly triangular postmental is in straight contact with the mental, in broad contact with the first and second infralabials, flanked by the elongate elements of the first, and in narrow contact with the central element of the second postgenial row. The malars are roughly as wide as long, in medial contact with the third infra- labials and broadly contact the second infralabials. The malars flank the two rows of postgenials, the interrupted first containing two and the second containing three to five segments. The second row of postgenials tends to be irregular and its segments may fuse with those of the first body annulus. There is no postmalar row, nor are the immediate postmalar segments enlarged by fusion. Dorsally the first body annulus includes one to two small seg- ments posterior to the third supralabial, the temporal, the postocu- lar, and the frontals. The dorsal segments of the nuchal region are enlarged and the corresponding ventral segments reduced. The dor- sal segments along the trunk consistently decrease slightly in size from the nuchal region towards the cloacal zone and onto the tail. There are 230-235 body annuli from the back of the third infrala- bial up to and including the pore-bearing precloacal. Irregularity of the segmentation or dorsal half-annuli are rare. A midbody annulus contains 14 to 16 (generally 16) dorsal and 16 to 18 (generally 18) ventral segments. The cloacal region is characterized by four large, circular preclo- acal pores, six to eight precloacal segments, 12 to 17 postcloacal segments and three to five lateral rows. There are 17 to 18 caudal annuli up to the smooth caudal cap, with the autotomy annulus fal- ling on the fifth to seventh postcloacal annulus. No specimens show autotomized tails or healed scars. Segments bearing rectangularly pigmented centers extend further ventrally on the autotomy annulus than on other caudal annuli. The tail is horizontally oval in section at its base, then reaches a vertically oval section, with an indented ventral surface in the terminal third. For the last three annuli it shows a sharp bilateral flattening so that it terminates in a blunt vertical keel. The lateral sulci begin after approximately the 45th body annulus and continue up to the level of the cloaca. These sulci are poorly de- fined and narrower than one bordering segment. There is no obvious dorsal sulcus. Lateral and ventral sulcal regions are characterized by slight indentations of the trunk. GANS & MATHER: AMPHISBAENIANS 33 14} om) eet oe = om 12+ fe) es | 10 o 10 15 20 Snout-vent length in cm Fic. 8. Amphisbaena medemi. Scatter diagram of tail lengths vs. snout-vent lengths for the available specimens. The middorsal segments of the midbody annulus are approxi- mately two times as long as wide. The ventral segments are approx- imately 1% times as wide as long. Range. — Known from the type locality only. Ecology. —The animals were taken during the construction of the fish cultural station of Inderena at the Laguna de Amajahuevo (De- partamento Atlantico) which lies on the shore of the Canal del Dique close to the village San Cristobal (Departamento Bolivar). The vil- lage is 1% hours by boat from the town of Soplaviento (Departa- mento Bolivar; 53km. from Cartagena) and lies between the lagoon (about 1% hectares) and the canal. All specimens, as well as some Leptotyphlops and Bachia, were collected at depths of approxi- mately 60 cm. to 1 m. in ditches easily excavated for foundations in 8I-ZI = =— XK yeso)y ‘ped xog 9I-ZI a1quliea 18-83 — 1€Z-002 8 x cate GO ¢ 9 orm) do 02-91 6I-LI 6 é L¥Z-EhS ‘pey espe i é rhe ae? v do “que xog 28-08 9 02 602-S0Z ‘yup “yup =~ —-XX = ([RAo)p yreqd yreqd 9I-FI 9I-F1 L-9 GS-1Z ShZ-F2S 9 = =x v ‘ped xog ST 9I-FI L-G SI-LT GEZ-08Z G x = Teao)y “peg xoq SI-9T SI-91 L 02-8T GS-81S y 4 = v do xog LI-¥1 FI-ZI euou PI-ZI 1SZ-98Z = a> v ‘peg ‘do ‘yup 9I-FI 91-21 L-G 91-21 VEZ-SZS xX =e v “peg ‘yup SI-9T 91-81 S-L ZS-8T L0Z-Z61 — = v do xoq ST-91 9I-FI euou 9-21 8SZ-6E% x = (9)F do xoq 02-FT SI-SI1 8-G SI-SI 9€Z-F1Z g = eo (6)8-F ‘jun ‘yu, 02-F1 9I-Z1 L-G Zo-LT €FZ-60Z x = v do ‘jyuy GZ-3S 9I-ST G 6I-9T 022-L02 Xx oy oh v do ‘ju ¥Z-0Z ST-91 9-S 12-61 123-661 x =e ¥(8) do ‘jug ¥E-1Z ST-9I auou 21-01 GZZ-L0Z Z xX v ao xog 02 ial auou 61-81 802-002 Z = bop & 4 (9)¥ do ‘yup ZG-81 LI-¥1 euou GI-O1 612-981 I — cea ¢ 9-F do xoq SI-9T PI-ZI euou SI-FI 8hZ-9ZS I — =X (9'S)F do ‘juny SI-FI 9T-ZI 6-9 vI-O1 81Z-66I 789381G ss SABRI ;SMOY selog Be (0) (oY) OJON ~sfeiqjueA = STBSIOG] aug TTnuuy TTnuuy jetoeds yetue4 [e13U9 A [esiog Apoqgptyl Auojojny jepned Apog munozuva 1asaulaqs 1az04 S$y19D18 1uapeu yjaLinds DD1zSauaf Dlax PpruUyos uayog DIaD9 1UUDUL 110Qa] “a toss1odky ‘3 sisuaanuos ‘3 SyDpNDod "1 suazouut ‘2 14NOQADQ *d pupqna ‘a BOLIOUIY YINOG U19Yy}IOU SSO1NB PUP SaTIJUY 94} SSO1NB YSva 07 JSAM DUaDgs1ydwY jo Satdeds [[eUIS 10} Seye4S Ia}OVIeY Jo AreuIUING *Z a[quy, 34 GANS & MATHER: AMPHISBAENIANS 35 the dry and sandy soil. No rain fell during the period of collection. Generally December is the end of the rainy season at this locality. The site of the excavation was unshaded by large trees, but cov- ered by pasture and scrubs. The mean temperature of the region is about 25-35° C. in the shade. Locals indicate that the general region sometimes floods during the rainy season. Etymology. —Named after Dr. Federico Medem M.., who collected the specimens, in recognition of his contributions to the biology of South American reptiles. Dr. Medem informs us that the animals are locally referred to as ‘‘tatacoas’’ or ‘‘ciegas.”’ DISCUSSION Table 2 (based upon the reports of Gans, 1962b, 1963, 1964; Gans and Alexander, 1962; Hoogmoed, 1973; Sponga and Gans, 1971; Thomas, 1965, 1966) summarizes the character states of the various small amphisbaenians of northern South America and the Antilles. It shows that the character states of Amphisbaena medemi are very close to those of A. fenestrata in body annuli, number of segments to a mid-body annulus, rostro-frontal contact, and number of preclo- acal pores. They differ in number of caudal annuli, in the tendency of A. fenestrata to have a medial sulcus at least posteriorly, as well as a postmalar row, and of A. medemi to have a longer tail and caudal autonomy. A. fenestrata shows a checkerboard pigmentation rather than the gradual fading of pigmentation that characterizes medemi. While the similarity to A. fenestrata is considerable, it is only an expression of a general similarity between such forms as A. xera and A. caeca, on the one hand, and of A. medemi and A. gra- cilis, on the other. A. medemi is similar to the Antillean form as it is the only form of the northern mainland that has fewer dorsal than Key to Table 2 opposite: 1. Key to abbreviations: Box, rectangular pigmentation in center of each seg- ment; CB, checkerboard dropping out of pigment; Fad. pigment fading and grad- ually from dorsal to ventral along sides of trunk; Unif., uniform pigmentation across entire surface of segments; X, first, second, and third genial or malar row present. 2. Key to Special States column. 1, Ocular fused to second supralabial; 2, extra dorsal half annuli; 3, rostral and nasals fused; 4, rostral and frontals in contact; 5, caudal tuberculation; 6, rostral and frontals generally in contact; 7, body seg- ments tuberculate; 8, mental and post mental fused. 36 FIELDIANA: ZOOLOGY, VOLUME 72 ventral segments to a midbody annulus. The various kinds of fu- sions (listed under special states) would seem to represent indepen- dent events rather than a sequence. We must again echo Sponga and Gans (1971) in deferring conclu- sions until additional and preferably internal character states of the new form have been compared with those of other species. This should presumably proceed in parallel with such analysis of the small amphisbaenians now assigned to other genera. The new form and theresults of some other recent papers (Sponga and Gans, 1971; Vanzolini, 1971; Stimson, 1972; Gans and Diefen- bach, 1972; Hoogmoed, 1973; Gans, 1974) permit amplification of the key to the American Amphisbaenia (Gans and Diefenbach, 1970). KEY TO THE AMERICAN AMPHISBAENIA [The following key to the Recent amphisbaenians of the Americas attempts to characterize the forms presently recognized on the basis of external characteristics. No effort has been expended to make the key natural but the most obvious and clearly apparent characteris- tics have always been emphasized. As certain characteristics such as pigmentation may occasionally be absent, and many species demonstrate caudal autotomy, we have avoided use of characteris- tics involving these and have provided multiple criteria for as many couplets as possible in order to facilitate identification. The key omits Amphisbaena polygrammica as being insufficiently charac- terized to permit diagnosis. ] Characterization of American Amphisbaenia: Body cylindrical, sometimes a ten- dency toward slight dorsoventral or lateral compression. Temporal musculature may be swollen and followed by a slight nuchal constriction in adults. Head often not differentiated from body in juveniles. Anterior third of body often slightly thick- ened but rarely with any significant diametric reduction between head and vent. Tail always less than 20 percent of body length. Skin annulated with dorsal and ven- tral half-annuli-‘meeting at, and sometimes continuous across, a lateral groove on each side which starts in the first third of the body and runs to the vent. Head may be rounded, pointed, spatulate or keel-shaped with or without keratin- ization. Mouth depressed, edges of lip inserted within head. Eye small, beneath skin, clearly visible in fresh specimens but with pigments sometimes bleaching out in preservative. Precloacal pores in a single row present or absent with their expression often ex- hibiting sexual dimorphism. Tail often with autotomy constriction at which it may break and heal into a conical stump without regeneration. Distal tip of original tail occasionally with keels, nobs or other modifications. GANS & MATHER: AMPHISBAENIANS 37 Two well-developed forelimbs, but no hindlimbs (head large and bluntly rounded; an enlarged azygous prefrontal; discrete nasals, preoculars and supraoculars; dorsal and ventral half-annuli poorly aligned along lateral sulci, with dorsal number 50 per cent higher; precloacal pores in two series, WIDEIY SEDALALEG) 5 tooth attr sale fan cere te oe Cee ve (Bipes) 10 No external evidence of fore or hindlimbs. ..............cccccecceees 2 Snout shovel-shaped with a more or less obvious horizontal edge (shields of pectoral region often markedly irregular; no precloacal pores, tail short, generally lacking an autotomy plane)...................00- 3 Head rounded, pointed or vertically keeled ..................0 ee eeee 5 Rostral folded around center of horizontal edge; three pairs of regular paired shields, but no azygous ‘shields, follow it along midline of the head; tail very short and blunt but with well marked narrow autotomy COBSUPICHION: 5. Csco es i Rae A ae ee Aulura anomala Rostral excluded from center of horizontal edge; one or more azygous shields along midline on dorsal surface of head; tail lacking autotomy COTNISELICGION SS weer cc eesa rs eal ean, eutelter al a tel svaitay ok cre oPetaricse enna. e eueve seein 4 Tip of snout broadly rounded; nostrils in discrete nasals; at least three enlarged azygous shields on midline of head; caudal tip dorsoventrally flattened, its dorsal surface covered with multiple small tubercles. ..... Rhineura floridana Tip of snout narrowly rounded or pointed; nostrils in a fused rostro-nasal shield; caudal tip short and bluntly rounded without tuberculation ..... (Leposternon) 15 Prefrontal azygous (flanked laterally by a postnasal supraocular segment; a single row of medially interrupted precloacal pores; tail short, round- ed, lacking autotomy plane; pigmentation pattern involving spots and blotches, more or less independent of the segmental arrangement)...... (Cadea) 14 Prefrontals paired: sncitei ca x is, aeeeegers oes ees Sle ae tee rea Eee Rae 6 Head keel-shaped; rostral very much enlarged, and ascending over the face, keeping the nasals, prefrontals, and sometimes the frontals from contact (keel sometimes keratinized; distal tip of tail may bear vertical ridge; tail acking autotomy: PIANO) ox: iess 8 sw Gases cater hve RnR pees ete ee 7 Head pointed or rounded, otherwise slightly compressed laterally; rostrals not separating frontals and prefrontals ................. ce eee eens 9 Frontals in broad contact on midline; rostral longer than high with a blunt vertical keel (rarely keratinized); (two precloacal pores separated by a median hiatus and three supralabials; 259-281 body annuli)........... Mesobaena huebneri Frontals (and prefrontals) in point contact or separated by ascending process of rostral; rostral higher than long with a sharp vertical keel; (four precloacal pores in a continuous series or two supralabials)...... 8 Postmental larger than malar; three supralabials; 214-249 body annuli {no supernumeraries); four precloacal pores in an uninterrupted row ...... Anops kingi 38 10. IT. 12. 13. 14. 15. 16. FIELDIANA: ZOOLOGY, VOLUME 72 Postmental not larger than malar; two supralabials; 358-372 body annuli (including supernumeraries); two precloacal pores in a medially inter- FUPLOGINO Waser: cnscataoic cus ere tene oreas DN stereiere seis he tverenewetess Anops bilabialatus Head with marked vertical compression; rostral in broad contact or ex- cluded from contact with enlarged prefrontals; neither parietals nor occipital segments enlarged; two medially-separated short rows of pre- Cloacaliporesiajcsisercccne on ier Poe Pe TRS (Bronia) 10 Head generally rounded, pointed or only faintly compressed; rostral with- out or with only narrow median contact with prefrontals, unless they have fused with nasals; a variable number of round or oval precloacal pores in a single row of procloacal segments without a major hiatus .... (Amphisbaena) 20 Nasal segments do not contact each other; 213-229 body annuli; 18-21/18- ZIIMADODY SOPMeNtE sc 6 iclag ow esas a ch ees aloes Bronia brasiliana Nasal segments contact each other; 281 body annuli; 28/24 midbody seg- MRMG TGS oir Foner reltesahooh sizes eres epost sie cred enw ee etre ut clisl ns soy eh eater eulonte (ote as Bronia kraoh Six to eight precloacal pores; three to five digits per hand............ 12 Two wide precloacal pores; five digits perhand........... Bipes biporus Three digits per hand; 142 ventral and 169 dorsal half annuli; 32-35 dorsal and 30-38 ventral segments to a midbody annulus .... Bipes tridactylus Four to five digits per hand; more than 148 ventral (and near 200 dorsal half annuli; apparently lower numbers of segments to a midbody annu- DUS) ctacchass ate elensaee waar hatly cracas cpaan puaheea nes wea siea nicer Oat aR ener 13 Most specimens with fewer than 159 half annuli; always fewer than 201 dorsal Half annals s/:5.0c oads ae cu ncire sc cm poate Bipes c. canaliculatus Most specimens with more than 165 ventral half-annuli; always more than 215 dorsal half annuli ............ Bipes canaliculatus multiannulatus Body annuli 175-218 (counted on ventral surface), caudal annuli 10-14, segments in a midbody annulus 25-33, snout rounded, blunt .......... Cadea blanoides Body annuli 274-320 (counted on ventral surface), caudal annuli 12-17, segments in a midbody annulus 32-39, snout domed, laterally com- PLOSSO asic is ts syaya el sake roe tess au Secoye ete pes Cadea palirostrata Only two rows of enlarged scales visible in dorsal view atop the head (the first of these the rostro-nasal, the second the azygous which may or may not be followed by a fringe of very much smaller segments in the occipi- tal region); suture pattern simple; pectoral region covered with very large geometrically regular shields ................. 0. eee ee eeeee 16 Three or more rows of enlarged shields atop the head, suture pattern often complex; pectoral region complexly divided or not ................ 17 Rostro-nasal followed by an enormous azygous shield which covers nearly the entire dorsal surface of the head, narrowly fringed posteriorly by a single row of very much smaller segments; head shield strongly keratin- ized; mental and postmental discrete; 253-305 postpectoral annuli..... L. scutigerum Iles 18. 195 20. 21. 22. GANS & MATHER: AMPHISBAENIANS 39 Rostro-nasal followed by a very large azygous, flanked on each side by tri- angular prefrontals and small temporals; headshields not significantly keratinized; a single mental-postmental segment; 378 postpectoral SI ssc iata ate og hea Aa ghee elalaiehoruie si ed eh his Men wate L. octostegum A single, very small first supralabial followed by a large second and a small third supralabial ic 2 3.5 scecciess foe: see pitta vex doe 9 9 ks a Sec en Bs 18 A very large first supralabial followed by a smaller second supralabial. . 19 Azygous shield in contact with rostro-nasal; frontals and medial tempor- als much larger than the azygous, longer than wide and arranged in a characteristic linear grouping; infraocular absent; a small first followed by a large second infralabial; pectoral shields in a regular arrangement of large shields reminding of an hourglass; dorsal number of postpector- al annuli generally equal to ventral, rarely up to 10 greater than; adults of medium size (snout-vent length equals 205-432 mm.).. L. polystegum Azygous shield separated from contact with the rostro-nasal by a wide suture between prefrontals; frontals about as wide as long, not sig- nificantly larger than the azygous, temporals much smaller, infraocular generally present; a very large first infralabial; pectoral segments only slightly enlarged, rounded sometimes with irregular longitudinal fu- sions; dorsal number of postpectoral annuli always five to 25 higher than ventral number; adults very large (snout-vent length equals 285- LUO) ari 00 bal aera eres Whee ot SNA Ri acd cine etn tT Ar heater ae eer REE L. infraorbitale Azyous always distinctly delineated; pectoral region covered by three to four pairs of elongated regularly arranged shields which may fuse except for the midline; number of dorsal and ventral postpectoral annuli more or less equal; 242 to 265 postpectoralannuli............. L. wuchereri Azyous often fused in various ways with adjacent head shields; a medial pair of elongate pectoral shields with other modified, possibly enlarged shields radiating out from these anteriorly; always significantly (5-25) more dorsal than ventral postpectoral annuli with the increase greater in the second, third, and fourth fifths of the trunk (177-242 postpectoral ATVYVUDLT!) Pet ee seek teas a aoe ee rte er SIR Pree tein Dee Ne AT L. microcephalum Nasals entirely separated from medial contact by posterior tip of ros- |g: | agar Ree rer as Cree UPR te CE aLY Or ater Seka Apr MPER Mert rnd rik, 21 Nasala im contactat midline. ccs ain ee ee ot ee eee eae 22 Body darker brown dorsally, lighter ventrally by checkerboarding, tail faintly conical, pointed, with 12 to 14 annuli, lacking autotomy, malar TOW. PLESEN Gate ero oe chet Nya donee ceesieaes ater ac cheese Seeker A. fenestrata Body darker brown dorsally, lightened ventrally by reduction of seg- mental centers. Tail rounded, with slight distal pinching, with 17 to 18 caudal annuli and an autonomy annulus at the 5th to 7th; no malar POW. W ekerc Sour creeaaemece teaser Aerie tonenie taveue ce aval te Ar arora rae ge A. medemi With constant and characteristic fusion of head shields.............. 23 Without major fusions of head shields (rostral, nasals, prefrontals, fron- tals, supralabials and oculars always discrete)..................-. 26 40 23. 24. 25. 26. 27. 28. 29. FIELDIANA: ZOOLOGY, VOLUME 72 Mental and postmental fused (243-247 body annuli; 6 precloacal pores; 33-39 segments per midbody annulus; nasals larger than prefrontals; caudal autotomy near 9th annulus).................... A. stejnegeri Mental and postmental discrete i.:.005562:5056 50's os cde x do noe Read 24 Second supralabial and ocular fused; rostral and nasals discrete (199- 240 body annuli; 10-18 caudal annuli; 4-6 precloacal pores; 26-34 seg- ments per midbody annulus; no postmalar row) ........ (A. cubana) 25 Second supralabial and ocular discrete; rostral and nasals fused (209- 243 body annuli; 17-22 caudal annuli; 4-9 precloacal pores; no postmalar TO Wi) eee ea ea I ea He NONE BEE oe to pecarras fay tty AAS A. manni 199-218 body annuli; 10-14 caudal annuli, with caudal autotomy......... A. c. cubana 226-240 body annuli; 14-18 caudal annuli, no caudal autotomy .......... A. cubana barbouri Very large and thick (adults near 40-50 cm. in length, more than 1.5 cm. in diameter); more than 65 segments to a midbody annulus; with prom- inent temporal; tail of same diameter as the trunk; lacking caudal autotomy; terminal caudal annuli weakly delimited; 4-10 precloacal POPES: soe Se anes ciere chan vere mictensiatetevenetabenah nets sha aiets eksusuenavaraneeeteaele A. alba Medium to large, generally more slender; between 48 and 65 segments to a midbody annulus; head acutely pointed, to a spatulate rostral tip; caudal autotomy lacking or if present the anterior body half annuli meet at the middorsal line in a posteriorly open herringbone pattern (A. angustifrons) 27 Small to medium-sized, adults less than 35 cm. long, less than 1 cm. in diameter; fewer than 48 segments to a midbody annulus; lacking caudal autotomy (diagnosis for A. miringoera based on illustration in Van- COMBI OT Sie 5 set aos ea eee seth Hak Ss Ses wore a PIE 28 Small to large, generally more slender; fewer than 85 segments to a mid- body annulus; head variously rounded or pointed, anterior body annuli crossing middorsal line in a more or less straight line; caudal autotomy BUNUMS OUVIOUELY PPOSSNG s si5 cc Secon a ala Gace in 5 ike aterdae ares Som aly ahs 38 Adults 28 to 41 cm. in snout-vent length; with body diameter 11 mm. or greater; generally 205-218 body annuli; annuli of the postnuchal region cross the back at right angles; no caudal autotmy; 13-19 caudal annuli A. a. angustifrons Adults less than 30 cm. in snout-vent length; with body diameter 11 mm. or less; generally 233-282 (rarely 210) body annuli; lateral half-annuli of the nuchal region typically meet in a posteriorly open angle pre- senting a ‘‘herringbone”’ pattern; autotomy between 5th and 9th caudal annulus; 18-21 caudal annuli ............... A. angustifrons plumbea More than 238 body arinuli: : 4 3c.c:ay is aise e os Oo old aa case aetna Be 29 Fewer than 232 Body Qn oad paca ms Lee eae Bee ees 33 More than 40 segments per midbody annulus.......... A. o. occidentalis Fewer than 41 segments per midbody annulus ...................-. 30 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. GANS & MATHER: AMPHISBAENIANS 41 Fewer than 20 caudal annuli More than 20 (22-24) caudal annuli; (250-262 body annuli; 12-15 ventral segments to a midbody annulus; two precloacal pores) . .. A. miringoera 14-16 caudal annuli; tail round ended; body cylindrical; head not smaller than trunk; second supralabials by far larger than 1st and 8rd supra- TaGiplee) chp toate arok vaies ain on Ore ae a Aru mates aatemuds A. bakeri 17-19 caudal annuli; tail clearly vertically keeled; body widened after nu- chal region; head very small relative to the trunk; second supralabials not markedly larger than 1st and 38rd supralabials................. 32 250-266 body annuli; 16 ventral segments to a midbody annulus......... A. s. steindachneri 239-245 body annuli; generally more than 16 ventral segments to a mid- POGY AUTUUB s 2 og aan aud aie ee tne eT ke A. s. borelli Tail sharply conical, tip only one-half diameter of base; very distinct nuchal constriction (fewer than 193 body annuli); snout pointed....... A, ridleyi Tail cylindrical; without conspicuous nuchal constriction (185 or more body annuli); snout broad Tail with marked vertical keel distally (26-31 segments per midbody ANINUNUS US aye, 5 cee eho Ae a NRA Ae pee rece has A. bahiana Tail with round end (29-42 segments per midbody annulus)........... 35 Postmental markedly larger than mental; postmalar row absent ...... 36 Postmental shield around the same size as mental; postmalar row pre- SONG iesseyspcuctst oe yee oi eeava tape Taedeh core pe rae Nt et Senos A. g. gonavensis Two precloacal pores; 213-231 body annuli; second supralabial faintly larger CRMN First os Scluctss citi g bate oA Oe ee ee Aes A. dubia Four precloacal pores; 185-219 body annuli; second supralabial markedly lation CRAR L786: 6 aes eA oa Cea een hea ea en eaA 37 Second row of postgenials with four or more shields; 9-14 caudal annuli A. i. innocens Second row of postgenials with four shields; 18-19 caudalannuli ........ A. innocens caudalis Two supralabials and three or fewer infralabials................+.-. 39 More than two supralabials and more than two infralabials........... 40 204-211 body annuli; 22-26 segments per midbody annulus; postoculars larger than prefrontals but smaller than parietals; two infralabials; four round precloacal pores; autotomy at the fourth to sixth of 23-25 caudal annuli; uniform medium brown, with little counter shading A. slevini 200-231 body annuli; 24-34 segments per midbody annulus; postoculars smaller than parietals which are smaller than prefrontals; 1% infralabi- als; four oval precloacal pores; autotomy at the 7th to 14th of 28-31 caudal annuli; uniform dark brown dorsally, generally light ventrally ... A. vanzolinii 42 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. FIELDIANA: ZOOLOGY, VOLUME 72 224-248 body annuli: 28-32 segments per midbody annulus; postocular smaller than parietals which are smaller than prefrontals; 2% to three infralabials; four large oval cloacal pores, autotomy at 6th to 7th of 21-22 caudal annuli; uniformly very dark brown dorsally and ventrally A. gracilis Without DOOLIDRISE TOW (ols oecis as Sess ote OS) Oe ae oh ele this ded 41 POStHiAlaT TOW DICNONG 3c k kt oon tos he Oiloe te ae Re ee Tae 48 TWO PROClomCAal PHOTOS: 5 46:53 sive bak aie alia Fale cco bo ee ha oni ee a 42 FP OUF.OF TOOTS PTECORCAl POTS seas 5s sees hE Ee ee a oa ae we eS 45 ewer than 162 body anni: 30.5. o.oo ele eek hoc ie abt em ean ee 43 More Chall 202 DOGY BONIN 3655 soos ce he ee oko RRS oy ae alae ee 44 Fewer than 162 body annuli; 24-28 segments per midbody annulus....... A. neglecta More than 161 (179-181) body annuli; 20-24 segments per midbody an- PRIN ooo Gy ck eo aveiy SPORE CREA S OE ED MOUS EN CRS A. silvestrii 203-220 body annuli; precloacal pores large and oval; nasals as large as or larger than prefrontals; parietals as large as or larger than frontals; TID OC CALTOUNG 5 oes os oa hy cose aia eo wie a IO Mee eos A. mitchelli 240-265 body annuli; precloacal pores round or faintly oval; nasals smaller than prefrontals; parietals never larger than frontals; tip of tail with VGICAN OB to ee ide Se eas hv OS BAW OO ORR eae A. roberti Postmental shield markedly longer and of larger area than mental..... 46 Postmental shield faintly larger, same size as, or smaller than mental 47 225-234 body annuli; 10-16 caudal annuli; 5th to 7th caudal annulus narrowed into autotomy constriction; first supralabial markedly smaller Lie CoAT | a7 Reaper avon Pantani alr i ar arch apap ptr nota tee a gran terra rea A. xera 231-245 body annuli; 19-22 caudal annuli; 7th to 8th caudal annulus nar- rowed into autotomy constriction; first supralabial around the same size as third (total length of adults, around 12 cm.)........ A. carvalhoi Fewer than 219 body annuli; 16-19 caudal annuli; 28-36 segments per mid- body annulus; no distinct nuchal constriction; tail with pigmentation POL CORTLO8 LUN oe shia s oo ssa rear ea ve elas Coes A. pericensis 222-226 body annuli; 19-24 caudal annuli; 26 segments per midbody an- nulus; very distinct nuchal constriction; tail more darkly pigmented CHAN DOGS os os Dares Saisie s SUE ES TAA COTES ONG A. nigricauda More than 54 segments per midbody annulus; head clearly narrower than body (acutely poitel) «0.66 6.65 ois cn scene anae nes A. camura Fewer than 56 segments per midbody annulus; head not markedly smaller CSNY CHAIN cis cig dace ea ate a inine e. EO shee Sia TR Rate © Ae SsherTe Bieta 49 More than four precloacal pores: 6. <.65. 62.4 oso vicisie Bsc ses see sess 50 Fewer than five prociouical poreg!s5.5 ic eso os as os weed awe sven 53 More than 10 precloacal pores; head and body to sixth annulus lacking PURINA ira ioe ec aon wee CA Rieke aisie ee te Rene A. leucocephala 51. 52. 53. 54. 55. 56. 57. 58. 59. GANS & MATHER: AMPHISBAENIANS 43 Fewer than 11 precloacal pores; head with same pattern of pigmentation BU EOE si bare arate k5 6 av acaia.s IGTS Fie eibia's Re aMoNBiaLb alo ers Wea we aes aieiet eats 51 Nasals larger than prefrontals; conspicuous checkered pattern of colora- tion; third supralabial often split; suture between mental and first infralabial sometimes fused 7.10.60) (5455-6 +. 04 oe tyetetaigios oe = A. fuliginosa Nasals smaller than prefrontals; without conspicuous checkered pattern; without splitting of third supralabial; suture between mental and first infralabial distance 3:0.) cos eas eee Oe a ae ee Oe ee na on ee Ce 52 Parietals of same size or larger than frontals; no preocular shields; pig- mentation of anterior part of body segments only; tail distinctly seg- MNONCOC CO ID fis ciccs ooo hese IE ae acto ac NS TEC A. mertensi Parietals never larger than frontals; preocular shields (split off second supralabial) often present; temporal bulges; tail relatively blunt with- out distinct segmentation until tip (bulging temporal region; uniform Drown, Ventral {Aging ) si 65 ova cs 5ig-oe ce ees see es A. pretrei More than 260 body annuli.................. A. occidentalis townsendi Kower thai 26t-bogy Gaius oo .5 = n” joa’ wu = Zz => 73 1978-78 FIELDIANA, ZOOLOGY$CH 590.5Fl 71 tet teas Le re Wem se Abbr ares Weer asenien