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NovttatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2590 JANUARY 5, 1976

CARL GANS Three New Spade-Snouted Amphisbaenians from Angola (Amphisbaenia, Reptillia) ::-i mu f- i055C- -V--{XH 74..oa*fTt fX- X¢ NovitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2590, pp. 1-1 1, figs. 1-4, tables 1-3 January 5, 1976

Three New Spade- Snouted Amphisbaenians from Angola (Amphisbaenia, Reptilia)

CARL GANS'

ABSTRACT

Angola has a diverse herpetofauna that in- Dalophia. Two Monopeltis each from a single cludes seven species of amphisbaenians. Besides locality are here described as new, as is the most two of the generalized genus Zygaspis, there are widely ranging species of Dalophia reported from five species of the spade-snouted genus Mono- seven localities in Angola and one in Zambia. peltis, and three of the spade-snouted genus

INTRODUCTION The extensive territory of Angola is for her- Most of the specimens came from museum col- petologists one of the least known parts ofAfrica. lections; some were donated by Dr. J. Quartau This is particularly unfortunate because there are and Mr. John Visser. The analysis disclosed indications that it may be one of the most inter- specimens of Monopeltis anchietae (Bocage, esting areas of the continent. Angola is ecologi- 1873), M. capensis (Smith, 1848), M. vandery- cally diverse. Its central and coastal regions are in- sti (Witte, 1922), and Dalophia pistillum (Boett- termediate between the humid forests of Zaire ger, 1895)', as well as D. welwitschii Gray and the extreme deserts of Southwest Africa, 'I here resurrect the name Dalophia Gray whereas these central and coastal regions are (1865), most recently used by Loveridge (1941). delimited inland by the Central African high- The reasons for setting aside Tomuropeltis (Laurent, 1947), a name which I used in my lands. This suggests that the region has high checklist (Gans, 1967), are given in Broadley, potential for relict coastal, swamp, and plains Gans, and Visser, MS. They are based entirely populations. upon a reconsideration of the status and char- A revision of the spade-snouted amphisbae- acter pattern of the types of Dalophia welwit- nians schii Gray, which show the truncate tail and of southern Africa (Broadley, Gans, and tendency toward intemasal contact character- Visser, MS.) has permitted me to re-examine istic of the assemblage that I (1967) referred to specimens from about 17 localities (fig. 1). Tomuropeltis. 'Research Associate, Department of Herpetology, the American Museum of Natural History; Professor of Biology, The University of Michigan, Ann Arbor.

FIG. 1. Records for Dalophia and Monopeltis in Angola and environs. Solid star, M. luandae; solid squares, M. perplexus; open squares, other Monopeltis; solid circles, D. angolensis; bulls-eyes, D. welwitschii; open circles, other Dalophia. (1865). With the exception of the last form, all named. Since the main report is still incomplete, records represent peripheral populations ofother- the new species are here described in the hope wise wide-ranging species. The review led to the that additional specimens may come to hand. discovery that two interesting forms of Mono- Terminology in this report is the same as in peltis and what was clearly the most widely dis- earlier descriptions for this group (Gans and tributed Angolan species of Dalophia were un- Latifi, 197 1; Gans and Lehman, 1973; Gans and 1976 GANS: AMPHISBAENIANS 3

Broadley, 1974). Specimens derive from the than fewer than seven supernumerary dorsal half- following collections and I thank the curators annuli to the first 50 body annuli. It differs (names in parentheses) who made the loans further from M. scalper in lacking precloacal possible: pores. It is easily differentiated from M. guentheri and M. schoutedeni by absence of precloacal AMNH, the American Museum of Natural His- pores, from M. anchietae by the fusion of head tory (R. G. Zweifel) shields in adults, and from M. anchietae, M. ca- CG, Gans collection, Ann Arbor pensis, M. zambezensis, and M. leonhardi by hav- CZL, Centro de Zoologia, Ministerio do Ultra- ing 15 to 16 rather than fewer than 11 caudal mar, Lisbon (J. F. L. Nunes, J. V. dos Santos, annuli. Its low number (fewer than 230) of body and Maria M. L. de C. Roque e Pinheiro) annuli distinguishes it from M. sphenorhynchus LCFM, Musee de la Chaux de Fonds, Switzer- mauricei, which has more than 275. land (J. L. Perret) Description. A small to medium-sized (203 to MD, Museo du Dundo Laboratoria de Biologia, Dundo, Lunda, Angola (these specimens 385 mm. snout-vent length) species ofMonopeltis were not examined) with the (apparently faded) dorsal surface of the NMB, Naturhistorisches Museum, Basel, Swit- trunk and tail speckled with dark pigment that zerland (K. Kramer) fuses into an accentuated area on the dorsal sur- NMSR, National Museum of (Southern) Rho- face of the tail. The species has 223 to 227 body desia; collection now at Umtali Museum, and 15 to 16 caudal annuli (up to the oval tip Rhodesia (D. G. Broadley) that shows some lateral compression), 29 to 36 USNM, National Museum of Natural History, (generally 30 or 32) dorsal and 14 to 20 ventral Smithsonian Institution, (J. A. Peters and segments to a midbody annulus, and four first George Zug) plus five to eight second postgenial segments. I am also grateful to the collectors for gifts of The azygous head shields are discrete at 218 mm. specimens, to Dr. Donald G. Broadley for hos- snout-vent length and show a broad isthmus of pitality and counsel, to the staff of the Centro keratin at 385 mm., but with significant lateral de Zoologia for information on the ecology of notches in the heavily keratinized shield (fig. 2). Dalophia angolensis, to Ms. Jean Barks for tech- There are no preoculars. The posterior processes nical assistance and to the National Science Foun- of the nasals (which segments neither touch each dation for support of these studies. Ms. Peggy other nor the lip) are excluded from contact with Hees drew the animals. the ocular by a wide contact zone between the third supralabial and head shield. The pectoral Monopeltis luandae, new species region is wide and very short. The medial pair of shields are by far the largest. They are widest Holotype. AMNH 111338, a male (subadult posteriorly and are joined anteriorly by a second or out of breeding season), collected during Feb- and smaller medial pair (derivative of the first ruary, 1973, by Dr. J. A. Quartau on the ground pectoral annulus) that widens anteriorly, with in Luanda on the road toward the mouth of the three to five pairs of much smaller shields clus- Quanza River, Angola. tering around the junction. One of the two of Paratypes. CG 5219, a smaller subadult male these may be replaced by a roughly parallel- collected on January 21, 1971, by J. A. Quartau sided lateral pair flanking the medial shields along at the airport, Luanda, Angola, and USNM their length. The lateral sulci are poorly ex- 20037 and 20038, respectively an adult male pressed and the dorsal ones are absent. Both and a very immature female collected during zones show major and crossing diagonal folding 1892 by Heli Chatelain, at "Loanda," Angola. lines. Supernumerary dorsal half-annuli number Diagnosis. Monopeltis luandae has a higher 14 to 19 in the first 50 (14 to 24 in the first 100) number (29 to 36) of dorsal segments to a midbody annuli and very few thereafter. The dorsal body annulus than does either M. scalper (13 to interannular sutures cross the trunk at right 19) or M. vanderysti (18 to 23) and differs from angles to its long axis. There are no precloacal M. vanderysti in lacking contact between nasals pores. Autotomy occurs at the third to fourth and preoculars and in having 14 to 19 rather caudal annulus. 4 AMERICAN MUSEUM NOVITATES NO. 2590

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FIG. 2. Monopeltis luandae. Dorsal (top), lateral (middle), and ventral (bottom) views of anterior portion of holotype, CG 5232. Etymology. The specific name refers to Lu- 5219 (near airport after rain); AMNH 111338 anda, the type locality. (road to mouth of Quanza River); USNM 20037, Locality Records. ANGOLA: Luanda: CG 20038. 00 00 WI 90 00

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Monopeltis perplexus, new species It is formed of six main shields, partial fusion occurring azygously between the lateral pairs, Holotype. AMNH 47732, a male taken by the and the almost triangular medial pair being by Vemay-Angola expedition at "Hanha or Cape- far the largest. Sutures radiate from the front and longo," Angola, between May 3 and July 20, a pair of enlarged segments of the prepectoral 1925. annulus inserts lateral to the medial pair. Lateral Paratypes. 3 males (AMNH 47731, 47733, and middorsal sulci are clearly expressed. Fewer 47734) and one female (AMNH 47735), part of than three supernumerary dorsal half-annuli oc- the same series. cur, mainly in the first and second 50 body an- Diagnosis. Monopeltis perplexus differs from nuli. The dorsal interannular sutures cross the M. sphenorhynchus mauricei in the greater num- trunk at right angles to its long axis. There are no ber of caudal annuli (22 to 24 rather than 8 to precloacal pores. Autotomy occurs at the fifth 12) and the clearly marked autotomy site; from to seventh caudal annulus. M. anchietae, M. leonhardi, and M. capensis in Etymology. The specific name characterizes the higher number of body annuli (more than the uncertainty of affinities and type locality of 261 rather than fewer than 221); and from these this series. as well as M. zambezensis by the more slender Locality Records. ANGOLA: Hanha or Cape- body shape and elongate tail with clear caudal longo: AMNH 47731-47735. autotomy, which these lack. It differs from the forms M. scalper, M. luandae, and M. vanderysti Dalophia angolensis, new species by the higher number of body (more than 261 rather than fewer than 245) and caudal (more Holotype. CZL 167 collected during 1959 at than 21 rather than fewer than 19) annuli. It Calombe, 7 km. "west" of Vila Luso, on Vila has fewer supernumerary dorsal half-annuli Luso-Moxico Road, Angola. (below 3 as contrasted with above 10) than any Paratypes. AMNH 111339; CZL 50, 204, 265, of the last group, except forM. vanderysti, from 292-300, 317, 318, 387, 388, 399, 400, part of which it is easily distinguished by its broad fusion the same series. of the head shields. Diagnosis. The new species of Dalophia can Description. A small (215- to 300-mm. snout- immediately be separated from the wide-ranging vent length in adults), slender species ofMono- D. pistillum, as well as D. gigantea (cf. Gans, peltis with the dorsal surface darkened by a 1967, for the status of these names), D. longi- speckling of individual melanophores evenly cauda, and D. luluae (new combination), by the spaced across the segments (mainly after the presence of caudal autotomy. Adults are much first third of the body and to just ventral of the shorter (shorter than 362 mm. rather than lateral sulci, but onto the ventral surface of the longer than 365 mm.) and stouter than D. tail). The speckles fuse into a darker tone on the ellenbergeri, and D. angolensis also has a much anterior part of the dorsal surface of the tail. shorter tail (approximately 40 mm., rather than The species has 261 to 270 body and 22 to 24 80 mm. in adults), and fewer caudal annuli (20 caudal annuli (up to the smoothly and paraboli- to 27, rather than 30 to 43). It differs from D. cally rounded tip), 12 to 17 (generally 16) dorsal welwitschii by having a greater number of body and 14 to 16 ventral segments to a midbody annuli (302 to 324, rather than 264 to 270) annulus, and four postgenial plus five postmalar and more regular contact of the two nasals, as segments. The azygous head shields are broadly well as nasalocular contact and angling of the fused and only a short, blind suture remains interannular sutures along the dorsal surface of anterior to the ocular at the base of a notch in the anterior trunk and of the tail. The species can the heavy keratinous shield (fig. 3). There are no in each case be further defined by supplementary preoculars. The slender posterior processes of characteristics. the nasals (which segments do not contact each Description. A slender, medium-sized (290- to other medially, but may touch the lip) are ex- 362-mm. snout-vent length) species of Dalophia cluded from touching the ocular by a wide con- that lacks pigmentation and has 302 to 324 body tact zone of the third supralabial and head shield. and 20 to 27 caudal annuli, 16 to 24 (generally The pectoral region is relatively short and wide. 18) dorsal and 12 to 18 (generally 14) ventral 1 976 GANS: AMPHISBAENIANS 7

FIG. 3. Monopeltis perplexus. Dorsal (top), lateral (middle), and ventral (bottom) views of anterior portion of AMNH 47732. segments to a midbody annulus and four or five second postgenials. The head shields are broadly first plus eight to 11 (generally eight or nine) fused and only the lateral blind sutures remain 8 AMERICAN MUSEUM NOVITATES NO. 2590

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(fig. 4). There are no preoculars, and the nasals All the CZL specimens were apparently col- (in narrow contact medially) almost always ex- lected within an area of about one hectare that tend long wings laterally to touch the oculars. was being cleared of vegetation and the very There are six elongate pectoral shields, the lateral sandy soil dug up to a depth of 15 to 20 cm. ones often asymmetrically subdivided and the They were taken in slightly moist sandy soil two medial pairs generally extending anteriorly covered with low scrub and isolated trees, rep- into one or two annular rows. Lateral sulci are resenting secondary growth over an area that had clearly expressed, but the dorsal sulcus is only formerly been planted. Approximately equal indicated by intersegmental alignment. The sixth numbers of Dalophia angolensis and Zygaspis or seventh caudal annulus bears the well-defined niger Broadley and Gans (1969) were taken and autotomy constriction. There are few super- the collectors did not note ecological differ- numerary dorsal half-annuli but the second 50 ences. The latter species does overlap the geo- body annuli tend to show irregular dorsoventral graphic ranges of D. angolensis and D. ellen- alignments. The intersegmental sutures of the bergeri. See Nunes and Tordo (1960) for a first 100 or so body annuli form anteriorly acute description of the region. angles with the midline. The middorsal caudal segments posterior to the autotomy constriction are twice or more as wide as those near the base LITERATURE CITED of the tail; the caudal interannular sutures form Bocage, J. V. Barbosa du an anteriorly acute angle on the middorsal surface. 1873. Reptiles nouveaux de l'interieur de Locality Records. ANGOLA: - (Monard, Mossamedes. Jor. Sci. Lisboa, vol. 4, 19371); LCFM (unnumbered); LCFM 848.46. pp. 247-253. Alto Cuilo: MD 5344 (Laurent, 1964). Cazombo: Boettger, Oskar MD 5790 (Laurent, 1964). Between Vila Luso 1895. Zwei neue Reptilien vom Sambesi. and Zool. Anz., vol. 18, pp. 62-63. Moxico, Calombe, 7 km. "west" of Vila Broadley, Donald G., Carl Gans, and John Visser Luso: AMNH 111339; CZL 50, 167 (specimen [MS.] Studies on amphisbaenians (Amphisbae- and two eggs) 204, 265, 292-300, 317, 318, nia, Reptilia). 6. The genera Monopel- 387-388, 399, 430. Lac Calundo: MD 5601, tis and Dalophia in southern Africa. 5705, 5744 (Laurent, 1964). Kakindo (Caquin- Gans, Carl do): LCFM (3 unnumbered); NMB 13332; 1967. A checklist of Recent amphisbaenians (Monard, 1931, 1937; Loveridge, 1941). Mu- (Amphisbaenia, Reptilia). Bull. Amer. panda2: NMB 13333 (Monard, 1937; Love- Mus. Nat. Hist., vol. 135, art. 2, pp. ridge, 1941). 61-106. ZAMBIA: Gans, Carl, and Donald G. Broadley Zambezi River (13001'S, 22044'E): 1974. A new dwarfed species of Monopeltis NMSR 2826. from the middle Zambezi Valley (Rep- Etymology. The specific name refers to the tilia: Amphisbaenia). Arnoldia (Rho- country. desia), vol. 6, no. 35, pp. 1-5. Biological Miscellanea. The Alto Cuilo speci- Gans, Carl, and Mahmood Latifi men had been caught by a hen (Laurent, 1964). 1971. Redescription and geographic variation of Monopeltis guentheri Boulenger (Amphisbaenia, Reptilia). Amer. Mus. 'Monard (1937) referred to seven specimens from Novitates, no. 2464, pp. 1-21. Kakindo on the Kuvangu River and Kuvangu Mission, Gans, Carl, and Grace C. Lehman identified first as M. ellenbergeri and then as M. granti 1973. Studies on amphisbaenians (Amphisbae- transvaalensis. Unfortunately he did not number his ma- terial nor are most of the LCFM specimens numbered. nia: Reptilia). 5. The species of Mono- Nevertheless it appears that those "no locality" speci- peltis from north of the river Zaire. mens do derive from Kakindo (=Caquindo). Occas. Papers, Mus. Zool., Univ. Mich., 2This locality was not mentioned for this form no. 669, pp. 1-34. (Monard, 1937). It may represent a cataloging error Gray, John Edward as does NMB 13330 the paratype of Amphisbaena am- 1865. A revision of the genera and species of buellensis Monard, 1931, which is entered as "Kuvangu" amphisbaenians with the descriptions from which locality Monard did not have specimens. of some new species now in the collec- 1976 GANS: AMPHISBAENIANS 11

tion of the British Museum. Proc. Zool. Bull. Soc. Neuchatel Sci. Nat., vol. 55, Soc. London, pp. 442-445. pp. 89-1 1 1. Laurent, Raymond F. 1937. Contribution 'a l'herpetologie d'Angola. 1947. Note sur les Amphisbaenidae d'Afrique. Arq. Mus. Bocage, Lisbon, vol. 8, pp. Rev. Zool. Bot. Africaine, vol. 40, no. 19-153. 1, pp. 52-63. Nunes, J. F. R., and G. C. Tordo 1964. Reptiles et amphibiens de l'Angola. 1960. Prospecq6es e ensaios experimentais (Troisibme contribution.) Publ. Cult. Cia apfcolas em Angola. Junta Inv. Ultra- Diamant Angola, no. 67, pp. 11-165. mar, Estud., Ensaios Doc., vol. 70, pp. Loveridge, Arthur 37-186. 1941. Revision of the African lizards of the Smith, Andrew family Amphisbaenidae. Bull. Mus. 1848. Illustrations of the zoology of South Comp. Zool., vol. 87, no. 5, pp. 353- Africa. London, pp. 1-28, pls. 1-78. 451. Witte, Gaston-Fran9ois de Monard, Albert 1922. Description de reptiles nouveaux du 1931. Mission scientifique Suisse dans l'An- Congo Belge. Rev. Zool. Africaine, gola. Resultats scientifiques. Reptiles. vol. 10, no. 1, pp. 66-71.

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