JMl -'*='^^—/" X^ ■-''-> /^^ v'Vf-. HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology JM UNIVERSITY OF KANSAS miscellaneous MUSEUM OF NATURAL HISTORY ^ \l^5^ MUS. COMP. ZOOL. LIBRARY JUiN 8 1973 hAHVARD. Systematics and Evolution of the Andean Lizard G^nus Pholidobolus (Sauria: Teiidae) By Richard R. Montanucci UNIVERSITY OF KANSAS LAWRENCE 1973 Ma/ 14, 1973 i UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly published in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publications, Museum of Natural History, began with nimiber 1 in 1946. Longer research papers are pub- lished in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subjected to critical review by intra- and extramural specialists; final acceptance is at the discretion of the pubHcations committee. Institutional libraries interested in exchanging pubUcations may obtain the Occa- sional Papers and Miscellaneous Publications by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas 66044. Individuals may purchase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Law- rence, Kansas 66044. University of Kansas Museum of Natural History Miscellaneous Publication No. 59 May 14, 1973 Systematics and Evolution o£ the Andean Lizard Genus Pholidobolus (Sauria: Teiidae) By Richard R. Montanucci Research Fellow Museum of Natural History University of Katisas Lawrence, Ka7%sas 66044 University of Kansas Lawrence 1973 University of Kansas Publications, Museum of Natural History Editor: Linda Trueb Managing Editor: William E. Duellman Miscellaneous Publication No. 59 pp. 1-52; 8 figures; 1 plate Published May 14, 1973 Museum of Natural History Unwersity of Kansas La\vrence, Kansas 66044 U.S.A. Printed By University of Kansas Printing Service Lawrence, Kansas CONTENTS INTRODUCTION 4 Acknowledgments 5 Materials and Methods 6 SYSTEMATICS 10 DELIMITATION OF SPECIES 10 GEOGRAPHIC VARIATION 16 REPRODUCTION AND BEHAVIOR 19 DISTRIBUTION AND ECOLOGY 20 EVOLUTIONARY DIVERSIFICATION IN PHOLIDOBOLUS 24 ACCOUNTS OF SPECIES 31 Pholidobohis Peters, 1862 31 KEY TO THE SPECIES OF PHOLIDOBOLUS 31 Pholidoholus affinis ( Peters ) 31 Pholidoholus prefwntalis new species 33 Pholidoholus montium ( Peters ) 34 Pholidoholus machnjdei new species 35 Pholidoholus annectens (Parker) 37 SUMMARY 38 RESUMEN 39 APPENDIX A: SUMMARY OF STATISTICS 42 APPENDIX B: SPECIMENS EXAMINED 50 LITERATURE CITED 52 INTRODUCTION Currently about 28 genera of small teiid lizards in Central and South Amer- ica are referred to the informal category of "microteiids." Pholidobolus is a genus of small, skink-like lizards which are less than 190 mm in total length. These liz- ards are characterized by relatively short, well-developed limbs, imbricate dorsal and ventral scales, and a color pattern composed of various shades of brown and cream longitudinal stripes. Populations of Pholidobolus are pres- ently known only from the Andes and interandean valleys in Ecuador, occur- ring principally between elevations of 1800 and 4000 m. These Hzards occupy a variety of habitats including semiarid grassland characteristic of the intermon- tane basins, montane forest on slopes and wet paramo above timber Hne. Dif- ferences in color pattern and morphol- ogy prevail between allopatric and para- patric populations. The current status of microteiid tax- onomy is chaotic. Considerable confu- sion has resulted from the definition of genera by single "key" characters, or combinations thereof. Furthermore, some "generic" characters, fundamental to microteiid taxonomy, are clearly con- vergent and parallel in their evolutionary history. Consequently, natural groups have been split on the one hand, whereas artificial, polyphyletic genera have been recognized on the other. Heretofore, Plwlidobolus has been composed of a single species, P. montium (Peters, 1862). However, it has become evident that with a systematic review of Pholi- dobolus, the taxonomic status of several other taxa should be reassessed. These taxa include Aspidolaemus affinis Peters, 1862; Macropholidus annectens Parker, 1930; Macropholidus ruthveni Noble, 1921; and Pholidobolus anomalus Miil- ler, 1923. Although Aspidolaemus affinis has been distinguished from Pholidobolus montium by the absence of femoral pores in males and the presence of prefrontal scales, I consider the taxa to be con- generic. Examination of additional ma- terial (including two new species) indi- cates that prefrontal scales and femoral pores are not useful generic characters in this group. The occurrence of pre- frontals varies inter- and intrapopula- tionally. For example, among popula- tions of a new species from central and southern Ecuador, the prefrontals occur at varying frequencies; whereas, another taxon, similar to P. montium, exhibits prefrontals in 99 percent of the material examined. Prefrontals are variably pres- ent or absent among individuals of some local populations of affinis. Contrary to reports in the literature, femoral pores are present in some males of affinis, to- tally absent in P. montium, and variably present or totally absent among males of some populations of one of the new species. The differences that do prevail between Pholidobolus and Aspidolaemus center principally on color pattern and relatively high scale counts, and are con- sidered to be of specific rank only. Be- cause Pholidobolus precedes Aspidolae- mus as a subgeneric name in Peters (1862) and because it was given generic status by Boulenger (1885), I place As- pidolaemus in the synonymy of Pholi- dobolus. Parker (1930) considered Macro- pholidus annectens to be morphologi- cally intenuediate between Pholidobo- lus montium and Macropholidus ruth- veni Noble. Rather than create another monotypic genus, Parker tentatively placed annectens in Macropholidus. The characters shared by annectens and ruth- veni include a clear lower eyelid disc, presence of uninterrupted rows of scales around the body, and absence of lateral body folds. Macropholidus ruthveni dif- fers from annectens in having smooth dorsals and two medial rows of greatly enlarged, hexagonal dorsal scales that are about twice as wide as long. By MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS contiast, the dorsal scutellation of an- nectens is like that of Pholidoholus mon- tium, showing weak keels or striations, and all being subequal in size. Further- more, additional samples of annectens show that the lower eyelid disc is pig- mented to varying degrees in some specimens and that the scale rows iiiound the body may be interrupted laterally by occasional scales of reduced size. In some Fholidohohis the lateral granules are reduced to a single row or entirely absent. Thus, the affinities of annectens seem much closer to Pholi- doholus than to Macropholidus. Crea- tion of a new genus to accommodate annectens seems unwarranted; I there- fore place it in PhoUdohohis. Provision- ally, I retain Macropholidus ruthveni generically distinct from Pholidoholus because none of the members of Pholi- doholus approaches ruthveni in the na- ture of the medial dorsals. A species from Cuzco, Peru, de- scribed as Pholidoholus anomalus (Miil- ler, 1923) has been rediscovered by Thomas H. Fritts. I have examined the specimens of anomalus and concur with Fritts' conclusion that the species does not belong in Pholidoholus. Proper allo- cation of anomalus will be the subject of a paper now in preparation by Fritts. Species treated in this systematic and evolutionary review include Pholidoho- lus affinis, P. montium, P. annectens, and two new species described herein. De- termination of the number of species to be allocated to Pholidoholus is based upon a discriminant function analysis of phenetically distinct populations in sym- patry, and a study of concordant varia- tion among different character states in the populations. Intraspecific geographic \'ariation is assessed for each species by means of Simultaneous Test Procedure (STP). This aspect of the study also includes a correlation analysis designed to demonstrate the relationships between climatic parameters, such as tempera- ture and precipitation, and morphologi- cal variation. The morphological, eco- logical, and geographical data thus ac- crued are employed in a Quantitative Phyletic analysis in an attempt to infer the evolutionaiy relationships within Pholidoholus. Largely as an outgrowth of problems encountered in the course of this study, I attempt to describe some methodological issues involved in the use of Quantitative Phyletic analysis. And finally, I test the use of quantitative phyletics as a means of identifying spe- cies. This is accomplished through a comparison of the population clusters delimited by Quantitative Phyletic analy- sis with those defined by discriminant function analysis and the criterion of character concordance. Acknowledgments The following curators kindly lent me specimens during the course of this study: Miss Alice G. C. Grandison, British Museum (Natural History) (BMNH); Dr. Walter Hellmich, Zoo- logisches Sammlung Miinchen (ZSM) via Dr. Thomas M. Uzzell; Dr. M. S. Hoogmoed, Rijksmuseum van Natuur- lijke Historic, Leiden (RMNH); Dr. Alan E. Leviton, California Academy of Sciences (CAS); Mr. Hymen Marx, Field Museum of Natural History (FMNH); Dr. Charles W. Myers, Amer- ican Museum of Natural History (AMNH); Dr. Giinther Peters, Zoolo- gisches Museum Berlin (ZMB); Dr. James A. Peters, National Museum of Natural History (USNM); Mrs. Doro- thy M. Smith, University of Illinois Mu- seum of Natural History (UIMNH); Dr. Charles F. Walker, University of Michi- gan Museum of Zoology (UMMZ); Dr. Ernest E. Williams, Museum of Com- parative Zoology (MCZ). Dr. William E. Duellman kindly permitted me to use the facilities and collections in the Mu- seum of Natural History, University of Kansas (KU). Most of the specimens collected during my study have been deposited at this institution. Field work in Ecuador was sup- ported by a Doctoral Dissertation Grant from the National Science Foundation 6 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY (GB-29567). A research assistantship from NSF was provided through the University of Kansas Committee on Sys- tematics and Evolutionary Biology dur- ing the 1971-72 academic year. This study could not ha\e been completed without the above support. Drs. William E. Duellman, Richard F. Johnston, Charles D. Michener and Linda Trueb provided editorial com- ments and suggestions which improved the quality of this work. Thanks also go to Thomas H. Fritts who willingly dis- cussed and criticized portions of this paper. Special appreciation is extended to Dr. Arnold G. Kluge, University of Michigan Museum of Zoology, who en- couraged and guided my learning of the methods of quantitative phyletics. Fi- nally, I am grateful to Drs. Bruce Mac- Bryde, formerly of Universidad Catolica del Ecuador, Quito, and Arthur C. Ech- ternacht, Boston University, for their companionship and help in the field. The Sums of Squares Simultaneous Test Procedure, Discriminant Function Analysis, and Quantitative Phyletic Anal- ysis were performed at the University of Kansas Computation Center. The Quantitative Phyletic Analysis was first performed on a preliminary data set at the University of Michigan Computation Center. Materials and Methods Materials. — This study is based on examination of 706 preserved specimens of PhoUdoholus. Of these, 609 speci- mens were used in detailed morphologi- cal study or statistical analysis. Samples for statistical purposes were drawn from specific, restricted localities, and repre- sent localized, presumably interbreeding, populations. In a few instances adjacent localities were pooled in order to in- crease sample size. The local samples used for statistical studies are listed in table 1. The locations of 46 place names mentioned in the text and tables are shown on the map of the Andean region of Ecuador (Fig. 1). Fig. 1. Andean region of Ecuador showdng the location of place names pertinent to this study. The localities are: Alausi (33), Aloag (11), Ambato (16), Banos (21), Biblian (36), Caja- bamba (28), Canar (34), Cayambe (4), Cerro Bueran (35), Cerro Llanganate (17), Cerro Picliincha (6), Chambo Grande (20), Chillo- gallo (10), Cotalo (23), Cubijies (25), Cuenca (37), Cumbe (40), Cutcliil (39), Giron (41), Gualaceo (38), Guamote (31), Guaranda (26), Ibarra (2), Lago Cuicocha (1), Latacunga (14), Licto (29), Lloa (9), Loja (45), Los Andes (18), Machachi (12), Mulalo (13), Ota- valo (3), Patate (19), Quito (7), Riobamba (27), Rio Chiche (8), Saraguro (43), San Antonio (5), San Jose del Chimbo (30), San Juan (24), San Lucas (44), San Miguelito "(15), Tixan (32), Urbina (22), Urdaneta (42), Zamora (46). Areas of 3000 m elevation and above are shaded. Methods. — Observations on the ecol- ogy and behavior of Fholkloholus were made during May and June 1971. The microhabitat preferences of the lizards as well as the general vegetative and cdaphic characteristics of some 30 lo- calities were recorded. Field observa- tions on behavior and reproduction in the lizards were made whenever possi- ble. About 80 lizards were brought to MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS the laboratory for detailed observations on beha\ ior. Color pattern variation in life was recorded from more than 300 animals. The recognition and delimitation of species herein is based on tsvo objective criteria: 1) geographical concordance among several independent character states; 2) sympatiic occurrence (sensti stricfo) of phenetically distinct entities, \\hich on the basis of life history data cannot be attributed to polymorphism. Character concordance is the coordi- nated disti-ibution of several different character states through geographic Table 1. List of local samples of Pholidobolus used in statistical analyses. The samples are designated alphabetically from north to south within the geographic range of each species. The abbreviation for each species is as follo\\'s: P. affinis (a), annectens (n), macbnjdei (c), montitim (m), prefrontalis (p). Locality Sampl a Size Code 2.9 km E Pan American Hwy, nr. Mulalo . 38 Aa 3 km SSW San Miguelito 6 Ba Ambato .. 19 Ca 2-3 km S Los Andes 11 Da 8.9 km SSE Patate 16 Ea Chambo Grande, 17.8 km W Baiios - - - 30 Fa Baiios - — 18 Ga 13 km E Riobamba 6 Ha 0.3-3.3 km S Ti.xan 14 la Loja Valley .. 26 An 12 km SW Cajabamba 7 Ac 14.2 km N Biblian 23 Be 9.1 km N Biblian __._„ 21 Cc 8-12 km S Cutchil 20 Dc 13-1.5 km E Loja 20 Ec Ibarra .. ._. _ .. 23 Am Ota\alo ._ .. ._ .. .. 54 Bm Cayambe 52 Cm 4 km W San Antonio 20 Dm San Antonio 17 Em Quito 21 Fm 2.5 km E Rio Chiche 31 Gm 2.9 km E Pan American Hwy, nr. Mulalo 8 Hm Guaranda .. . _. _ 31 Ap 3.3-4.9 km S Tixan 29 Bp Alausi ___ 16 Cp 0.5 km S Rio Caiiar on Pan American Hwy 9 Dp Cutchil 12 Ep 4 km E and 6 km N Cuenca 11 Fp space. Concordance was analyzed by plotting character states for each char- acter on maps of Ecuador. Points of equal expression of a character were connected by lines (isophenes) which enclosed geographic regions. Areas of maximum overlap among states of dif- ferent characters could be revealed by superimposing isophene distributions for all characters. Such zones of concord- ance (core areas) mark the codistribu- tion of character states of different char- acters and serve to delimit phenetic entities to which specific names were applied. The analysis of concordance was based on nine characters (six mor- phological, three color pattern), chosen because of the ease of defining their character states. Independence among characters was tested by correlation analysis where possible. Series of specimens taken from zones of sympatry were identified initially to species on the bases of head scutellation and color pattern. Although morphologi- cal intermediates were not apparent, hybrids conceivably could resemble one of the parental species in both sets of characters while favoring the other in meristic traits. To test for maintenance of integrity. Discriminant Function Anal- ysis was applied to meristic characters only, and the results were used to cor- roborate the preliminary identifications. The parental species were represented by reference samples taken from areas geographically removed from the zone of sympatry. The reference localities were chosen on the basis of gross eco- logical and climatological similarity with the area of contact, thereby increasing the validity of comparisons among sam- ples. A mixed sample from the zone of sympatry was treated as an unknown (test sample). Meristic characters were differentially weighted relative to their within- and between-groups variability using variance-covariance mathematics. A discriminant multiplier was calculated for each character, and this was multi- plied by the value of its respective char- acter. All such values were summed for 8 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY each individual to yield its discriminant score. The scores of the reference sam- ples and test sample were then plotted on frequency histograms and compared. Sums of Squares Simultaneous Test Procedure (Power, 1970) was employed as an aid in describing major trends of variation among meristic characters within species. However, the analysis was not used to support the recognition or delimitation of species. As will be discussed later, the meristic characters in PhoUdoboltis are responsive to a com- plex of environmental parameters and are not very useful for diagnostic pur- poses. Therefore, statistical similarity between tsvo populations need not be equated to close phylogenetic relation- ships. The STP analysis employed a test of significance after an overall analysis of variance demonstrated significant differ- ences among means. Samples were ranked in decreasing order of their means, and sums of squares were calcu- lated by sequentially adding means (starting with the largest) until a maxi- mal non-significant subset was delimited. The procedure was repeated, each time deleting one or more of the largest means in the previously described subset until another non-significant subset was de- fined, and until all samples were in- cluded in at least one such subset. A significance level of .05 was used for statistical tests. The STP tables (Tables 9-23) comprise Appendix A. Quantitative Phyletic Analysis (Kluge and Farris, 1969; Kluge, 1969) was used to infer evolutionary relationships among the species of PhoUdoboltis. The pro- gram generates a most parsimonious dendrogram depicting the evolutionary relationships of the OTUs (Operational Taxonomic Units) in terms of cladistic events and amount of character change (patristic distance). The major strong points of the methodology include an objective technique for weighting char- acters according to their variability within and between OTUs, and an op- erational means of employing Wagner's ( 1961 ) criteria for the estimation of primitive character states. In this study, the input OTUs were represented by the most objective units — local samples of PhoUdoboltis, that were analyzed without regard to tax- onomic status. Thus, the manner in which local samples clustered and the relative patristic distances between clus- ters could be used in the evaluation of the species recognized by the criteria of character concordance and sympatry. The weighting function employed in the analysis was computed by dividing the standard delation of the sample means of the ith character by the aver- age standard deviation of the ith char- acter. The value obtained was then multiplied by the mean of the respec- tive character for each sample. Char- acters were thus accorded weight pro- portional to their conservatism. More- over, the weighting function corrects for the scaling effect shown in biological variables — the larger the structure, the greater is its variability (Kluge and Ker- foot, 1971). Averaging the standard de- viation can be statistically hazardous. However, I know of no better way to obtain a weighting function based on the amount of within- and between-group variability. Applying a separate char- acter weighting in each species of PhoU- doboltis would not compensate for po- tential error since there is usually as much variation within a species (local populations) as between species (see Appendix A). For discussions on the concept of conservatism and its applica- tion to evolutionarv taxonomy see Farris (1966) and also' Fisher (1930) and Guthrie (1965). Proponents of Quantitative Phyletic Anah'sis ha\'e stressed the merits of the methodology but have almost totally ne- glected its shortcomings. By enumerat- ing certain problems I hope not to dis- courage prospective users, but rather to instill caution and direct their efforts toward improving the methods. When working with taxonomic cate- gories above the species level, it is possi- MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 9 ble for a given character to be conserva- tive in one group of organisms, yet rather variable in another. The weight- ing function is necessarily an average N'alue, and to apply an average weight for the character to both groups is less than a satisfactory practice. Another problem is that cladistic results can be influenced by the relative number of low- and high-weighted characters. The low weights of several variable char- acters in combination may outweigh a conservative one. If considerable homo- plasy is associated with the variable characters, misplacement of OTUs may occur; this can also result from the presence of correlated characters in the data set. Such characters also will un- duly augment the patristic distance be- tween OTUs. Finally, caution is urged concerning interpretation of the amount of evolutionary change depicted on the phyletic consh'uct. Patristic distance, expressed by the relative lengths of the lines connecting OTUs, is obtained by summing the individual differences be- tsveen the means of each unit character. For example, assume two OTUs, A and B, have mean values of 2.1, 2.4, 2.6, and 2.5, 2.7, 2.8 for three characters respec- tively. The patristic distance between A and B is 0.9, the sum of 0.4, 0.3, and 0.2. However, the difference between each pair of means may not be statis- tically significant; each pair of values may represent the same parametric mean. Because patristic difference is not based solely on significantly different mean values, it is possible to accrue a considerable difference between two OTUs (especially with application of a weighting function) when, in reality, a small difference, or no difference at all exists. Therefore, it is not justifiable to name a taxon on the basis of patristic distance alone. Characters. — Specimens of Pholido- holiis were described by a set of 21 ex- ternal, meristic characters and one mor- phometric character. Most of the char- acters were utilized in the Quantitative Phyletic Analysis and Simultaneous Test Procedure. In order to minimize experi- mental error, only specimens in a good state of preservation were studied in de- tail. Correlation analyses were per- formed to test for redundancy among characters. Deletions of certain charac- ters will be explained in later portions of this paper. Each character was ex- amined for sexual dimorphism. Only femoral pores and head diameter were found to be significantly different be- tween the sexes (females lack femoral pores and have relatively slender heads ) ; for these characters sample means are based on males only. Osteological comparisons ( x-ray; cleared and stained specimens) were made between the species of Pholidobo- lus (N=64). Slight differences in the relative positions of cranial elements were noted among specimens, but this variation was individual in nature and does not serve to distinguish species. Characters of the axial skeleton were found to be highly conservative, defining PhoUdoholus as a group ( see Accounts of Species), but not useful at the species level. For these reasons I was unable to utilize osteological data in my evo- lutionary analysis of the members of PhoUdoholus. William Presch (pers. com.) currently is comparing the oste- ology of the micro teiids. I therefore have made no attempt to place Pholi- dohohis in any relationsliip to other gen- era. Furthermore, its apparently close relatives, Pantodactyhis and Prionodac- tijhis (Uzzell, 1969), are poorly known taxonomically; thus, an analysis of inter- generic relationships is prematiu"e at this time. Counts were taken between struc- tures whose homologues could be readily identified in each individual among the different samples. For example, count- ing the number of "occipitals" would necessarily demand the ability to dis- tinguish these scales from "temporals." In some lizards identification would be difficult and uncertain, and counts would be subject to error. However, counting the scales in a row between the postero- 10 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY lateral edges of the orbits obviates the need for discrimination and insures high repeatabiHty. Definitions for each character studied are provided below: Number of prefrontals (NPF). — Tri- angular scales (usually a pair when present) situated between the anterior corners of the ocular orbits and the posterior margin of the frontonasal plate. Scales between orbits ( SBO ) . — Scales along a straight line between the postero- lateral corners of the ocular orbits. Number of supraoculars (NSO). — Scales between the posterior and an- terior margins of both orbits. Number of scales along margin of upper jaio (SUJ). — Scales between the anterior edge of the external auditory meatus and the anterior-most extreme of the snout on one side only. Number of scales along margin of lower jaw (SLJ). — Scales between the anterior edge of the external auditory meatus and the anterior-most extreme of the lower jaw on one side only. Number of gular and jaw scales ( SG}). — Scales along a straight line from the collar fold (well developed fold im- mediately anterior to shoulders) to the anterior-most extreme of the lower jaw. Number of ventrals (SGV). — Scales along a straight line from a point medial and just posterior to collar fold to the anterior margin of the vent. Number of dorsals (DEL). — Scales in a straight hue from the anterodorsal edge of the external auditory meatus to a point immediately posterior to the in- sertion of the hindlimb. Number of temporals ( NTS ) . — Scales along a straight line between the pos- terior corner of the orbit and the anterior edge of the external auditory meatus. Number of scales around the body (SAB). — Scales around the body at a point 13 scale rows posterior to the collar fold. Number of scales around the tail (SAT). — Scales around the tail at a point 9 scale rows posterior to the vent. Number of scales along the forelimb ( SAP ) . — Scales dorsally along a line be- tween the insertion of the limb and the tip of the fourth digit. Number of supradigital scales on fin- gers.— Scales along the dorsal surface of the third finger (SF3) and fifth finger (SF5) from their insertions on the palm of the hand to the tips of the digits. Number of supradigital scales on toes. — Scales along the dorsal surface of the third toe (ST3), fourth toe (ST4), and fifth toe (ST5) from their insertions on the sole of the foot to the tips of the digits. Number of femoral pores (FP). — Number of scales bearing femoral pores on both thighs and expressed as a total. Lateral granules ( LG ) . — Granular scales separating ventrals from the en- larged dorsals at a point at the 13th ventral scale row (coded 1); enlarged dorsal scales in contact with ventrals at that point (coded 0). Lower eyelid (LEL). — Lower eyelid transparent (coded 0) or pigmented (coded 1). Lower eyelid scales (LES). — Num- ber of distinctly enlarged scales covering the right lower eyelid. Head width (HW).— Widest point of the head (measured with dial calipers and recorded to an accuracy of 0.1 mm); the value is expressed as a proportion of the snout- vent length. Snout-vent length (SVL). — Distance between the anterior-most extreme of the lower jaw and the vent; measured with a metric rule to an accuracy of 1 mm; used only to express the head width character. SYSTEMATICS DELIMITATION OF SPECIES states and sympatry (sensu stricto) of I have applied the criteria of con- phenetically distinct populations in the cordance among independent character recognition of species of Pholidobolus. MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 11 Character concordance is the coordi- nated distiibiition of character states of several different characters through geo- graphic space. These areas of maximum character overlap or core areas (see Methods) correspond to phenetic species (sensii Sokal and Crovello, 1970). As explicitly shown by Sokal and Crovello (1970), tlie biological species concept defined by Mayr (1963) is non-opera- tional. I have made no attempt to define species of PhoUdobolus on the basis of their interbreeding properties; my infer- ences regarding reproductive isolation apply only to the local populations in- vestigated. However, the analysis of zones of contact and sympatry has facili- tated the delimitation of the species by providing data on the existence of phe- netic gaps or sharp phenetic transitions. Characier Concordance. — Nine char- acters and their respective states used in studying geographical concordance are: A) lateral body pattern — a. striped, b. ocellar, c. uniform; B) dorsolateral pale stripe — a. reaches snout, b. does not reach snout; C) lateral neck and tail stripes on males — a. both brown, b. both red, c. former brown, latter absent, d. both absent; D) number of supraoculars — a. two, b. three; E) number of pre- frontals— a. absent, b. always present (ca. 100%), c. usually present (ca. 75%); F) femoral pores of males — a. absent, b. rarely present (ca. < 26%), c. usually present (ca. 90%); G) texture of dorsals — a. striated-placoid, b. striated-keeled; H) lateral fold — a. present, b. absent; I) lower eyelid scales — a. three to six opaque scales, b. one clear or semi- opaque scale. For some of the aforementioned char- acters, certain states have been arbitrar- ily defined to facilitate presentation of the data, but this action does not weaken or change the general conclusions to be drawn from the information. The dis- tributions of individual states of the nine characters are shown in figure 2. Zones of maximum concordance (core areas) derived by superimposing the nine dis- tributions are shown in figure 2J. Popu- FiG. 2. The distributions of individual charac- ter states (enclosed by solid lines and desig- nated by lower case letters) of nine characters ( A-I ) . Zones of concordance ( J ) were obtained by superimposing the distributions of the nine characters. Names applicable to the core areas (J) are as follows: P. montium (a), P. affinis (b), P. annectens (c), P. prefrontalis (d), and P. tnacbrydei (e). Localities of sympatry or near contact between species of PhoUdobolus are Mulalo ( mu ) , Tixan ( ti ) , and Cutchil ( cu ) . A map like that shown in figure 1 was originally used to plot character state distributions. See text for discussion. lations in core area (a) conform in mor- phologic detail to the description of PhoUdobolus montium. Likewise, the names of P. affinis' and P. annectens are available for the populations in core areas (b) and (c) respectively. Two new names, PhoUdobolus prefrontalis and P. macbrydei, are proposed for the populations in core areas (d) and (e) respectively. Sympatry. — Where two species of PhoUdobolus come into contact, the zone 12 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY of overlap is extremely narrow. Sym- patric associations were studied at two localities (Fig. 2J). Near Mulalo (mu, Fig. 2J), a mLxed population of P. affinis and montium was found inhabiting a stone wall. The former species outnum- bered the latter about five to one. The two species are apparently tolerant of each other, for mixed basking congre- gations were observed. On the basis of head scutellation, there was no evidence of hybridization, although individuals of P. affinis were noticeably darker in color than usual, thereby superficially resem- bling P. montium. The superficial simi- larity may be attibuted to a common set of environmental pressures acting upon the populations. Discriminant function analysis (Fig. 3) based only on meristic characters (Table 2) substantiates the apparent lack of morphological intermediates. The population of P. affinis at Mulalo has converged towards the allopatiic popu- c D A B — ' 1 1 1 1 28 30 32 34 36 38 40 LINEAR DISCRIMINANT SCORES Fig. 3. Histograms of linear discriminant scores for samples of Pholidobolus. Allopatric reference samples are A. P. montium (n = 35, x = 31.5) from the Rio Chiche, and B. P. affinis (n = 35, x = 37.9) from Ambato and SSE Patate. Test samples C. (P. montium) and D. (P. affinis) are from a zone of sympatry near Mulalo. Each square represents a single specimen. See text for discussion. MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 13 Table 2. List of meristic characters and their discriminant multipliers used in Discriminant Function Analysis comparing Pliolidobohts af- finis with P. montiuin (A) and P. affinis witli P. prcfrontalis ( B ) . Characters Discriminant Multipliers A B SUT 0.079 0.614 SLJ 0.075 -0.206 SGJ 0.083 0.454 SGV 0.282 0.222 DEL 0.146 0.162 NTS 0.485 0.345 SAB 0.025 -0.075 SAT 0.268 0.272 SAF 0.172 0.060 SF3 0.224 0.241 SF5 -0.280 0.036 ST3 0.259 0.208 ST4 0.162 0.101 ST5 - 0.482 - 0.365 lation (sample A) of P. monthtm. The sample of P. montium from the zone of sympatiy seems to have undergone char- acter displacement; its discriminant scores fall at the lower extreme of values shown for the allopatric reference sam- ple of P. nwntiiim. The displacement could be attributed to competitive pres- sure from P. ofjinis. On the Pacific slopes of the Andes, populations of PholidohoJus offinis come in contact with those of P. prefrontalis. The two species were found to be sym- patiic in a limited area about 3.3 km south of Tixan (H, Fig. 2J). The zone of s\'mpatry was observed briefly on two occasions. Several instances of actual or attempted homospecific copulation be- tween lizards were seen, and adults in pairs were also noted to be conspecific. In captivity a mixed group of P. affinis and P. prefrontalis was maintained for several months. Sexual interactions ob- ser\'ed during that time were all homo- specific. Examination of a mixed sample from the zone of sympatiy did not provide evidence for hybridization on the basis of cephalic scutellation or color pattern. Discriminant function analysis based on meristic characters (Table 2) revealed two atypical specimens (Fig. 4). One of the specimens is identified as P. pre- frontalis (KU 141103) and the other as P. affinis (KU 140934). The two speci- mens are typical of their respective spe- cies in color pattern and cephalic scutel- lation, and also appear normal in gross gonadal morphology. However, both specimens show extreme values for most of the meristic characters used in the analysis, suggesting that the lizards may be hybrids. If so, interspecific matings are probably relatively infrequent, chance events, because only two lizards in a series of 42 specimens show inter- mediate discriminant scores. No other zones of contact were found between species of PholidohoJus. How- ever, on the east side of the Cuenca Valley south of Cutchil (cu, Fig. 2J), populations of P. prefrontalis and mac- hn/dei occur within 2 km (airline dis- tance) of each other, and are separated by a vertical distance of only 320 m. Sympatric association may be revealed by additional intensive searching. The adjacent populations show no evidence of hybridization and the hiatus between them is sufficiently narrow to indicate that they are distinct species. Interspecific Comparisons. — Among the characters most useful in distinguish- ing species of PhoUdohohis are those associated with color pattern, cephalic scutellation and dimensions, number of femoral pores, number and condition of the lower eyelid scales, and presence or absence of lateral body folds. Diagnoses of the species are deferred for a later section of this paper. However, a sum- mary of the morphological information is provided in table 3. Meristic charac- ters describing body and limb scutella- tion are not diagnostic because of the considerable overlap among the species in the ranges of variation. Statistical differences do exist among some inter- sDccific and inti-aspecific local samples. Specific comparisons can be made in only a \'ery general sense. Most samples of PhoUdohohis affinis show relatively high mean values compared to those of the other four species. Samples of P. 14 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 30 32 34 36 38 40 42 LINEAR DISCRIMINANT SCORES Fig. 4. Histograms of linear discriminant scores for samples of Pholidobolus. Allopatric reference samples are A. P. pre/rojitofo (n = 31, X = 32.6) from Guaranda, and B. P. affinis (n = 35, x = 41.9) from Ambato and SSE Patate. Test samples C. (P. prefrontalis) and D. (P. affinis, stippled) are from a zone of sympatry near Tixan. Each square represents a single specimen. See text for discussion. machnjdei and P. annectens have rela- tively low mean values, and samples of P. montium and P. prefrontalis have sam- ple means usually falling between the high and low extremes of most meristic characters (see Appendix A). Note that the information in table 3 consists of the range of variation for a given character and the range for the sample means. A mean for each species was not calculated (except for P. annec- tens) because it is of no value for com- paring local samples. The intraspeciiic local samples are sufficiently different from each other that the value of a mean of a species would be directly dependent on the relative proportion of individ- uals representing low and high sample means. Since the localized sample is an objective unit within which individuals are taken at random, the sample mean is most useful for comparati\'c purposes. Cephalic scutellation is helpful in identifying species of PhoUdohohis; how- ever atypical, individual variation not associated with geographical trends, oc- casionally occurs. The usual number of supraoculars in P. affinis is six (three on each side); four supraoculars are typically present in the other species. MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 15 Table 3. Comparison of morphological characters among tlie species of Pholidobolus. Range of variation and range of sample means ( in parenthesis ) are provided for each character. Character P. affinis P. montium p. prefrontalis p. machnjdei P. annectens NPF 0-3 0 0-3 0-3 0 (1.9-2.1) (0.0) (1.3-2.2) (0.0-1.5) (0.0) SBO 4 2-4 2-5 3-4 4 (4.0) (3.9-4.0) (3..5-4.0) (3.6-4.0) (4.0) NSO 5-8 4-6 4-5 4-6 4 (5.9-7.0) (4.0-4.2) (4.0-4.1) (4.0-4.5) (4.0) SUJ 11-15 10-14 9-14 9-13 11-12 (12.4-13.6) (11.0-12.4) (9.9-12.1) (10.7-12.1) (11.6) SLJ 11-15 9-14 10-13 9-15 10-13 (12.2-13.5) (10.4-12.2) (11.0-11.8) (10.4-11.9) (10.8) SGJ 16-26 15-22 15-20 13-22 1.5-20 (19.3-22.5) (17.4-19.4) (16.1-18.8) (1.5.6-18.8) (17.7) SGV 27-35 24-31 24-32 22-28 25-30 (29.3-32.1) (26.6-28.4) (26.8-30.1) (24.8-25.9) (27.3) DEL 45-55 35-50 37-46 31-43 40-48 (47.4-52.5) (40.7-46.2) (.39.5-45.3) (40.7-46.2) (42.6) NTS 5-10 4-8 5-9 4-8 5-8 (6.3-8.3) (5.3-6.6) (.5.3-7.1) (4.8-6.2) (6.5) SAB 34-54 31-45 31-45 27-41 23-29 (38.2-50.0) (35.0-42.1) (36..5-40.4) (.30.3-.37.0) (26.5) SAT 22-34 19-27 18-28 18-26 17-23 (24.7-30.5) (20.5-24.6) (19.9-24.3) (20.6-22.0) (19.1) SAF 22-29 18-27 20-26 18-25 20-25 (24.0-27.1) (21.8-24.6) (21.3-24.1) (19.6-23.1) (22.4) SF3 9-12 7-10 8-11 7-10 8-10 (9.8-10.8) (8.3-9.2) (8.1-9.8) (8.0-8.5) (8.1) SF5 5-8 5-8 5-7 5-7 5-7 (6.4-7.1) (6.0-6.4) (5..5-6.7) (5..5-6.3) (6.0) ST3 10-14 8-13 8-12 7-11 8-12 (11.1-12.3) (9.9-10.8) (9.5-10.6) (8.7-9.7) (10.0) ST4 12-17 10-16 10-15 9-14 11-15 (13.9-15.4) (12.2-13.2) (11.9-13.6) (10.6-12.4) (12.2) ST5 8-13 8-12 8-10 6-11 7-10 (9.3-11.1) (8.8-9.8) (8.2-9.3) (7.9-8.8) (8.3) LES 3-6 3-6 3-6 3-6 1-2 (3.3-4.1) (3.8-4.4) (3.6-4.4) (3.6-4.4) (1.0) T,FL 1 1 1 1 0-1 (1.0) (1.0) (1.0) (1.0) (0.1) LG 0-1 0-1 0-1 0-1 0 (0.5) (0.3-0.5) (0.4-0.5) (0.1-0.5) (0.0) HW 0.151-0.180 0.15.5-0.175 0.150-0.177 0.164-0.204 0.154-0.166 (0.157-0.167) (0.1.59-0.168) (0.L58-0.168) (0.168-0.195) (0.161) FP 0-4 0 0 0-11 0-2 (0.0-0.5) (0.0) (0.0) (0.0-10.3) (0.4) Some individuals of P. affinis taken near Los Andes, Patate, Chambo Grande, and Tixan, however, have as many as eight. The increase is usually due to fragmen- tation of the middle, triangular supra- ocular into a large inner and small outer scale. A few specimens of P. affinis from Baiios have five supraoculars ( three plus two); the two scales on one side are markedly different in size. Among the other species of PJiolidoboliis, more than four supraoculars have been noted in relatively few specimens. A single indi- vidual of P. prefrontalis from Alausi has five scales. In P. montium five or six supraoculars are present in a few speci- mens from Otavalo, Cayambe, Quito, and Rio Chiche. Some specimens of P. machnjdei from near San Juan, Caja- bamba, and Loja have five or six scales. The supernumerary supraoculars are usually small scales fragmented from the 16 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY anterior or posterior supraocular. How- ever, in one specimen of P. montiinn from Rio Chiche, a triangular scale is wedged between the anterior and pos- terior scales, identical to the arrange- ment found in P. affinis. Prefrontal scales are typically present in P. affinis and P. prefwntalis, absent or present in P. machrydei, and absent in P. montium and P. annectens. A few P. affinis from Baiios and San Miguelito lack one or both prefrontals. Some indi- viduals from Ambato and Tixan have three prefrontals; the exti-a one is a small, square scale lying between the two normally present. PhoUdoholus pre- fronfalis shows similar variation in the prefrontal scales. A few specimens from near Guaranda have three prefrontals, and two lizards among five taken at Biblian lack prefrontals entirely. The typical cephalic scutellation and variations in supraocular pattern among species- of PhoUdoholus are shown in figure 5. GEOGRAPHIC VARIATION Results of the Simultaneous Test Pro- cedure (STP) analysis (summarized in tables 9-23 in Appendix A) pro\'ide a basis for this discussion of intiaspecific geographic variation in Phohdohohis. Major trends are described for 15 mer- istic characters studied in the following number of local samples: P. affinis (9), P. annectens (1), P. machrydei (5), P. montium (8), and P. prefrontaUs (6). In tln-ee-fourths of the samples, the char- acters SGV and DEL are significantly correlated (0.05 level); characters SF3 and ST3 are also significantly correlated with each other and with all other digit counts in most samples. Intraspecific Patterns of Variation. — Among samples of PhoUdohohis affinis no latitudinal clinal trends are apparent. However, high mean values for most characters are present in geographically central samples, and there is a tendency for reduction in mean values among peripheral samples from the north, south- west and east. Among the nine samples, those with means in the highest third include Chambo Grande ( 13 charac- ters), Ambato (11 characters) and San Miguelito (six characters). Peripheral samples with means situated in the low- est third include Baiios (13 characters), Mulalo (10 characters), and Tixan (six characters). Eleven characters (SUJ, SLJ, SGJ, SGV, DEL, SAB, SAT, NTS, ST3, ST4, ST5) exhibit high mean values in two or more of the centrally located samples and low values in two or three peripheral populations. Characters SF3 and SAP, with high mean values in three central locations, show reduction in only one peripheral locality' (Baiios). Characters LES and SF5 with high mean values at two central locations show re- duction in the outlying samples from Mulalo and Bafios respectively. In PhoUdohohis affinis the texture of Fig. 5. Head scutellation in PhoUdoholus. A. P. affinis, KU 127129; B. P. prefrontaUs, KU 134912; C. P. mo7itium, KU 118086; D. P. machrydei, KU 121259; E. P. annectens, KU 121196. Atypi- cal supraocular patterns are A'. P. affinis, FMNH 28036; B'. P. prefrontaUs, KU 121238; C. P. mon- tium, KU 141261. MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 17 the dorsal scales varies from striated to keeled \\'ithin and between local popu- lations. The sample from Mulalo is dis- tincti\e in that nearly all indixiduals ha\ e smooth dorsals; weak striations are evident in a few specimens. Four characters appear to show clinal \'ariation in PlioJkloholus Dwchrydci; howexer, only five samples were large enough for analysis. Mean values from smaller, geogi'aphieally interspersed samples seem to support the apparent trend. Mean values of the counts show an increase from north to south with or \\athout slight reversals. The southern- most sample from east of Loja has the highest mean x'alues for 13 characters. Comparison of four specimens from 9.5 km south of Saraguro with the southern- most and northern samples reveals a step in the clinal patterns of two charac- ters. In the characters SAF and SAB the sample from near Saraguro is simi- lar to the northern samples, indicating a sharp step somewhere between the Saraguro area and the locality east of Loja. A more or less gradual clinal ti^end is shown by the characters ST4 and ST5. The sample from east of Loja is also unusual in having a high frequency of prefrontal scales (ca. 75%). In other samples of PhoJkJoholus machrydei, the prefrontals are either absent or have a low frequency. In a sample of 23 lizards from 14.2 km north of Biblian, only two individuals have prefrontal scales. Among 20 lizards from 8-12 km south of Cutchil, only one lizard was found with prefrontals. Femoral pores are variably present or absent in local samples of P. mac- brydei. Ranges and sample means are: four males from 10 km west of San Juan, 10-11 (10.3); ten males from 9.1 km north of Biblian, 2-11 (6.5); 15 males from 14.2 km north of Biblian, 1-11 (5.6); 11 males from east of Loja, 0-9 (3.3); five males from 12 km southwest of Cajabamba, 0-5 (2.6). Complete ab- sence of femoral pores is known only in the sample taken from 8-12 km south of Cutchil. Typically, the dorsal scales in P. 7nac- brydei are weakly striated or placoid. The latter condition is especially evi- dent in samples from wet paramo at high elevations. The sample from east of Loja is distinctive in having a rela- tively high proportion of individuals with striated or keeled dorsals. The only known specimen of P. macbrydei from the upper Jubones valley has keeled dorsal scales. The characters LES and SGV are statistically homogeneous among the five samples of P. macbrydei. Samples of FhoUdobolus prefrontalis show no general geographic trends, clinal or otherwise. However, for many char- acters, high sample means seem to be associated with xeric localities. For ex- ample, high mean values for 11 of the 15 characters (all except LES, DEL, ST5, NTS) are present in the sample from near Rio Caiiar which is an espe- ciallv xeric localitv. Means for five char- acters (SUJ, SLJ,'SAB, SAF, LES) show relatively high values in the sample from near Tixan, which also is noticeably dry. By conti'ast, in the more mesic situa- tions characteristic of the northern part of the Pacific drainage (Guaranda), the lower Andean slopes (Alausi), and the Cuenca Valley, the sample means for many characters have relatively low val- ues. Nine characters (SLJ, SGJ, SGV, DEL, NTS, SAB, SAF, SF5, LES) have low mean values in the sample from Guaranda; eight characters show low values in the sample from near Cuenca, and seven exhibit low means in the sample from Alausi. On the east side of the Cuenca Valley at Cutchil, a popu- lation of P. prefrontalis shows moderate values for most characters and low values for SLJ, SUJ, NTS, and LES. In PhoJidobohis monfium the only geographic trend evident is the presence of relatively low mean counts in samples from the northern and southern extremes of the geographical range and relatively high values in centrally located samples. At the northern end of the range, the population from near Otavalo shows low values for 11 characters (all except NTS, 18 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY SAT, SGJ, LES), and the sample from Ibarra has low mean values for SLJ, SGJ, SAB, SAT, SF3, SF5. In the south- ern sample from near Mulalo, seven characters show low mean values (SGJ, NTS, SAT, SUJ, SGV, DEL, SAF). Cen- tral samples, such as from San Antonio, 4 km west of San Antonio, and from near Rio Chiche in the Valle de los Chillos, show high mean values for 14, 11, and seven characters respectively. These central locations are noticeably dry. Valle de los Chillos, for example, sup- ports some species of plants knowTi only from dry areas to the south ( Bruce Mac- Bryde, pers. com.). Therefore, an ap- parent association betu'een dryness and high meristic values may apply to both P. montium and P. prefronialis. LES is the only statistically homogeneous char- acter in P. montimn. Geographic variation in P. annectens is not discussed herein because all lo- calities were pooled into a single sample. Clhnatic Parameters and Morpho- logical Variation. — In the foregoing dis- cussion I alluded to an apparent rela- tionship between aridity and meristic variation in Pholidoholus prefro7ifalis and P. montium. In an attempt to un- derstand the influence of enxironmental factors on character variation, simple correlation and regression analysis was applied to 20 morphological traits and two ecological parameters (rainfall and elevation) for 11 local samples of Pholi- dobolii.s. Correlations involving elevation were non-significant. However, inverse relationships were found between a six year average for annual precipitation and the following characters: lateral granules (r = ~ 0.797, P < .01); scales between orbits (r= — 0.707, P < .05); number of temporals (r = — 0.645, P < .05); number of prefrontals (r = — 0.640, P < .05). In PhoUdohohis montium and P. prefrontalis a weak positive correlation was found between precipitation and the number of dorsals ( r = 0.750, P < .06); lack of significance at the 0.05 level is attributed to small sample size ( N = 7 ) . Dorsals and femoral pores were the only characters showing posi- tive r values among the samples. In Pholidoholus machnjdei an apparent re- duction in number of body scales with cool-wet conditions may be adaptive. Lizards with relatively few, large body scales would ha\e a large surface to volume ratio, enhancing solar absorption in the cool, cloudy montane environ- ment. Furthermore, if heat loss occurs through the epidermis between scales (assuming the scales to be impermeable structures) the total area for potential heat escape is reduced by having a re- duced number of body scales. The nu- merous, small body scales which charac- terize some populations of P. affinis and xeric-adapted P. prefrontalis would en- able the lizard to adjust rapidly to clironic heat levels in warm-dry areas. That is to say, a small-scaled lizard once in the shade can lose heat faster than a large-scaled lizard. This hypothesis re- mains to be tested experimentally. For alternative models see Soule and Kerfoot (1972). Because rainfall data were not available for localities of P. machnjdei, the apparent relationship between num- ber of scales and precipitation could not be tested by regression. Ehrlich and Raven ( 1969 ) ad\anced the thesis that natural selection is at one and the same time, both the primary cohesive force and the primary disrup- tive force producing organic diversity. They argued that the influence of gene flow between two populations depends ultimately on selection. Thus, if two populations exist under markedly differ- ent selecti\'e regimes, they will diverge despite gene flow between them. Con- versely, two populations under the same or similar selective regimes will tend to remain similar even in the absence of gene exchange. Given that climatic variables charac- teristic of a locality' at least crudely represent the selective regime, then morphological similarity between two populations of Pholidoholus should pro- portionally reflect the climatic similarity' of their localities. Regression analysis MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 19 was used to test this hypothesis. Esti- mates of morphological similarity based on patiistic distance (amount of weight- ed character divergence ) was taken from 15 intraspecific pairs of local samples. Se\ eral measures of environmental simi- larity between localities were obtained, as follow: 1) an annual rainfall index calculated by summing mean monthly rainfall differences between localities; 2) an annual temperature index calcu- lated as abo^'e, using mean monthh' tem- peratures; 3) a climatic index including both temperature and rainfall as calcu- lated abo\e. In addition, geogi-aphical distance (adjusted for topography) was calculated between pairs of localities. Patristic distance was then regressed against the variables. Non-significant correlations were obtained for geo- graphic distance ( r = 0.064 ) , tempera- ture index (r = 0.289) and the climatic index (r= 0.054). However, the corre- lation between patristic distance and the precipitation index was significant for a six year annual average ( r = 0.527; P < .05) as well as for one year (r = 0.528; P< .05). The influence of environmental fac- tors on morphological variation in Pholi- dobohis is far from understood. The data indicate that precipitation is cor- related with at least five meristic scale characters and can be of predictive value when analyzing overall morphological similarity between populations. Cause and efi^ect relationships can be tested experimentally by incubating the eggs of PhoUdoholus under diff^erent tempera- ture and moisture regimes. Heritability studies could be used to partition varia- tion into its emironmental and genetic components. Temperature, although not correlated with meristic variation, prob- ably interacts with other factors in a complex manner. There is no relation- ship between geographic distance and morphological similarity between popu- lations of PhoUdoholus. Thus similarity cannot be attributed to gene flow. REPRODUCTION AND BEHAVIOR PhoUdoholus usually deposits two eggs at a time; occasionally a lizard lays a single egg on two consecutive days. Sites of egg deposition obviously are used by more than one individual, and/ or repeatedly by the same individuals. Multiple clutches are known for PhoU- doholus montmm, prefwntalis, and mac- hnjdei, and suspected for the other spe- cies. At Lago Cuicocha, one site con- tained 21 eggs of P. 7nontium. At the bottom of a rock pile near Cutchil about 12 eggs (including old, empty shells) belonging to P. prefrontalis were discov- ered. South of Cutchil, nest sites under flat rocks contained four to six eggs of PhoUdoholus machnjdei. Measurements from the eggs of PhoU- doholus (ranges followed by means in parenthesis) are as follows: 6.6-7.6 x 11.6-13.2 (7.14 X 12.38 mm) in nine eggs of P. montiwn; 6.2-9.0 x 12.0-15.5 (7.42 X 13.76 mm) in five eggs of P. prefron- talis; 5.7-7 A X 10.5-12.2 (6.46 x 11.62 mm) in ten eggs of P. machnjdei; 6.8- 7.1 X 12.3-12.8 (6.95 x 12.52 mm) in four eggs of P. annectens; 8.0 x 14.3-14.8 (8.0 X 14.55 mm) in two eggs of P. affinis. The snout-vent length of juveniles measured shortly after hatching are: 25 mm in one P. affinis; 22-23 mm in two P. annectens; 21 mm in one P. mac- hnjdei; 22-23 mm in two P. prefrontalis. Females of P. affinis with large, yolked follicles have Ijeen collected on 11 May and 21-23 June. Individuals with oviducal eggs have been taken on 26 May and 23 June. In P. prefrontalis, lizards with large, yolked follicles have been collected on 11 May, 9 June, and 13 July. Females containing oviducal eggs have been taken on 12-14, 16, 24 Mav, 9 June, 13, 15, 25 July. Females of P. montium with large, yolked follicles have been collected on 26 February and 16 June. Lizards with oviducal eggs have been taken on 16 June, 17, 25"july, 31 October. In P. machnjdei, large, yolked follicles have been found in lizards col- lected on 14 May, 18-19 June, and 30 July. Lizards with oviducal eggs have 20 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY been collected on 13-14 May, 10, 19 June. In P. annectens, lizards with large, yolked follicles have been collected on 13 and 21 June; oviducal eggs have been noted in females taken on 12 June. Whether the reproductive cycle in Pholidobolus is seasonal or continuous remains to be determined, because the above data simply reflect the field activ- ity of collectors. Collections tlu'oughout the year are needed for each species. Telford (1971) suggested that competi- tion between sympatric microteiid liz- ards in Panama is probably reduced in juveniles by non-overlapping reproduc- tive cycles and, perhaps, differences in diet. My data, although limited, do show that P. affinis, montium, and pre- frontaUs are reproductively active during the same months. Differences in diet could reduce competition between sym- patric species of PhoUdoholns, but this remains to be investigated. Actual or attempted copulation has been observed in all species of Pholi- dobolus. The following descriptions of receptive and non-receptive behavioral patterns are t\'pical for all members of the genus. Initially, the male crawls alongside the female, frequently licking her body. If receptive, the female re- mains still, permitting the male to grasp her nape with his jaws and to insert a hemipenis while his left or right hind leg straddles her lower back. If um'e- ceptive, the female crawls continually with her body slightly raised posteriorly and her tail writhing slowly. Frequently the female stops and makes several over- hand pawing motions with either fore- limb. Males exhibit this same unrecep- tive behavior if approached sexually by other males. There is no evidence that males are aggressively territorial. In the field, males were sometimes observed in close proximity, and encounters between foraging individuals went without inci- dent. Some microteiid lizards, for ex- ample Eiispondylus (Fouquette, 1968), show pronounced aggressive behavior. The mechanisms of species discrimi- nation and mate selection in PhoUdoho- lus remain to be investigated. The fol- lowing evidence suggests that olfactory cues may be principally involved: 1) sympatric species of Plwlidohohis appear to be reproducti\'ely active during the same months; 2) courtship behavioral patterns among the species are identical or at least very similar; 3) considerable licking activity occurs when members of the courting pair first encounter one another. DISTRIBUTION AND ECOLOGY The Andes are the dominating physi- cal feature of Ecuador, traversing the entire country from north to south. This highland region presents a formidable barrier to east-west dispersal. The Andes consist of two Cordilleras, joined in sev- eral places by east-west spurs or ridges which separate some ten intermontane basins. The Eastern Cordillera is com- posed principally of gneiss, mica, shist and other crystalline rocks. The Western Cordillera, however, is derived from porphyritic eruptive Mesozoic rocks and some Mesozoic sedimentary strata, chiefly Cretaceous (Sauer, 1965). Most of the intermontane basins or valleys are situated between the Eastern and Western Cordilleras. The northern- most basins are of complex surface form, having a terrace-like peripheral rim of high terrain, and a low, inner basin excavated by river action. The Ibarra Basin, for example, shows terrace rem- nants between 2133 m and 2438 m; far below at an elevation of 762 m, lies the flat valley bottom along the Rio Chota, a tributary of the Rio Mira. Similarly, the Quito Valley shows a volcanic ter- race along the lower slopes of Cerro Pichincha, ending as an abrupt scarp which drops to the floor of the basin some 2286 m above sea level. The Quito Basin is separated from the Central Val- ley to the south by a ridge of highlands connecting the two cordilleras. The Cen- tral Valley encompasses two basins. The northern Latacunga Basin is drained by headwaters of the Rio Pastaza; the MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 21 southern Riobaniba Basin also is drained by headwaters of the Rio Pastaza, which flo\\'s eastward to join the Rio Maranon. The Riobamba and Latacunga basins are partially separated by the massive base of Mt. Chiniborazo. On the Pacific slopes, the Alausi Basin is formed along a west-flowing tributary of the Rio Na- ranjal. To the southeast lies the Cuenca Valley, drained by headwaters of the Rio Santiago, a tributary of the Rio Mara- iion. Still farther south lie the smaller basins of Oiia and Loja. Populations of Pholidoholus occur throughout the Andean region. The dis- tributions of all of the species of Pholi- doholus are essentially allopatric or para- patric (Figs. 6 and 7). Pholidoholus tuontiuiu occupies the northern basins of Ibarra and Quito, extending at least as far south as the northern end of the Central Valley. Pholidoholus affinis in- habits the Central Valley, but reaches the Pacific slopes of the Western Cor- dillera near Tixan. Pholidoholus pre- frontolis occurs along the Pacific slopes from Guaranda south to Caiiar and in the adjacent Cuenca Basin. Pholidoho- lus machnjdei is distributed along the crests and slopes of the Western and Eastern Cordilleras as well as on the interconnecting highland ridges. All species of Pholidoholus inhabit the basin floors and adjacent slopes except P. mac- hnjdei, which occurs principally on the upper slopes and crests. Elevational dis- tributions (range followed by mean) for each species are as follows: P. ofinis, 1800-3050 (2611.1 m); P. montium, 2000- 3190 (2726.5 m); P. annectens, 2150- 2335 (2208.8 m), restricted to Loja Basin; P. machnjdei, 2315-3962 (3190.4 m); P. prefwntalis, 2400-2885 on Pacific slopes, and 2295-2620 in Cuenca Basin (2552.8 m). Pholidoholus machrydei is the most mesic-adapted member of the genus, in- habiting wet paramo grasslands and montane forests. In paramo and sub- paramo situations P. machnjdei may be found among clumps of Stipa, under rocks, or in Yucca or bromeliads. Pholi- doholus tnonthim and P. annectens are found in similar but less mesic situations. Pholidolwlus annectens is endemic to the Loja Basin, which has an average an- nual precipitation of 942.5 mm (Garcia, 1963-69). The average annual precipita- tion for localities inhabited by P. mon- tium varies from 1039.6 to 590.2 (821.2 mm); this suggests that the species has some tolerance for dryness. Pholidoho- lus prefwntalis is primarily a xeric- adapted species. Average annual pre- cipitation varies from 836.4 to 239.0 (481.1 mm) at localities inhabited by the species. Although P. prefrontalis in- habits areas that are generally drier than those occupied by P. montium, the tol- erance limits of the two species overlap. Pholidoholus affinis exhibits a broad moisture tolerance, occurring in areas with an average annual precipitation of only 2.39 mm to as great as 1367.5 mm (634.4 mm). Thus, P. affinis is consid- ered the most generalized member of the genus from the viewpoint of precipi- tation. All species of Pholidoholus receive solar heat through basking and substrate absorption. Basking sites include rocks, agave leaves, bromeliads, or clumps of bunch gi-ass. In the high, windswept paramo, P. machnjdei can be found sun- ning in protected situations near the bases of large bunch grasses. In sub- paramo areas, the bases of Yucca are used for basking sites. Pholidoholus machnjdei remains active even after in- solation has been blocked by clouds and fog. The lizards continue to receive heat through substrate absorption by pressing the body close to the surface of Yucca or other objects. By so doing, the lizards can remain abroad even after the am- bient temperature has dropped to 18° C. Pholidoholus usually occm"s in local- ized, high-density populations associated with rock piles, stone walls, or agave fence rows. Distributional localization is emphasized by man's clearing of the habitat and construction of rock piles and stone walls. Pholidoholus may be more sparsely dispersed in undisturbed 22 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 6. Distribution of Pholidobolus montium (triangles), P. affinis (dots), and P. annectens (square) in the Andean region of Ecuador. Crosses mark zones of sympatry between P. montium and P. affinis at Mulalo and Baiios (questionable). Areas of 3000 m elevation and above are shaded. MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 23 Fig. 7. Distribution of Pholidobolus machnjdei (dots) and P. prefrontalis (triangles) in the Andean region of Ecuador. Cross marks zone of sympatry behveen prefrontalis and affinis (see Fig. 6) near Tixan. Areas of 3000 m el e\ation and above are shaded. 24 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY habitat except where natural rock outcrops harbor locaHzed, dense pop- ulations. In undisturbed, dry paramo, individuals of FhoJidoholus ofinis are relatively more scattered than those as- sociated with stone fences. Usually a single individual is found near each clump of bunch grass or some other form of cover. In wet subparamo habitat, single males or male-female pairs of PlioUdohohis 7nacbrydei are found occu- pying individual clumps of Yucca that are scattered among the bunch gi'ass. By contrast, a dense population of P. mac- hnjdei was found in a rock pile near the edge of a pasture. Except for apparent differences in moisture tolerance, all species of PhoJi- doholiis are similar in their ecology and habits. Presumably, subtle ecological differences exist between geographic areas occupied by each species, and the superiority of each species in its respec- tive area prevents encroachment and displacement by another. Perhaps this explains why different species of Pholi- doholus do not occur sympatrically over broad regions. By contrast, the teiid genus Procfoponis is sympatiic with Phohdoholus montium and prefronfalis over extensive areas. Apparently the secretive habits of Procfoponis and per- haps other differences permit it to co- exist \M'th the free-foraging, basking PhoJidohohis. EVOLUTIONARY DIVERSIFICA- TION IN PHOLIDOBOLUS Selection of Ancestral Character States. — A controversial, yet necessary, procedure in evolutionaiy systematics is the inference of primitive character states. Wagner (1961) recognized sev- eral criteria for estimating primitive states. According to Wagner, an ances- tral state is likely to be 1) present in numerous representatives of closely re- lated gi'oups; 2) more widespread within a group than any one derived state, and 3) associated with states of other char- acters known to be primitive by other evidence. In determining the wide- spread nature of a character state one does not simply count numbers of taxa showing a particular state. A character state is widespread if it occurs in sev- eral taxa that otherwise ha\'e little in common. Thus, if a particular phyletic line were much more successful than others and produced many more species, etc., one would not erroneously consider a character state in that line to be primi- tive merely because many taxa showed that condition ( Kluge and Farris, 1969 ) . Kluge (1969:20) employed the Prim Network as an objective method of ap- plying Wagner's criteria. The network is an undirected, minimum length link- age system connecting OTUs on the basis of overall similarity. Wagner's criteria are founded on the premise that dixer- gent evolution is more probable than convergent evolution in any gioup. Kluge (1969) reasoned that the most primitive set of character states is that OTU situated at the cladistic center of the network. This OTU directly links the most OTUs and has the smallest mean and standard de\iation of interval lengths, and thus represents the point from which the largest number of differ- ent evolutionary directions can be real- ized. In order to adequately estimate the "widespread " nature of a character state (criterion 2) and avoid confusing an evolutionarily successful phyletic line as a primitive one, a Prim Network analysis must include numerous "distant" taxa. Ob\ iously use of the Prim Network has serious limitations in evolutionary stud- ies dealing with closely related species within a single genus. The rcliabilit}' of this approach to estimating primitive states increases as higher taxonomic cate- gories are selected for study. Teiid genera other than PhoJidohohis could not be included in my Prim Net- work analysis due to paucity of material for statistical manipulations. Therefore, another approach was used to estimate ancestral states. Each character was examined separately in PhoJidohohis and MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 25 representatives of 16 other loiid genera (actual specimens or literature descrip- tions). The relative primitiveness of in- di\idual character states was evaluated on the bases of ecological information and Wagner's criteria. The resulting com- posite suite of ancestral states was then compared with the actual OTUs of Pholi- (lo])ohis. For purposes of discussion, I ha\e grouped morphological characters, as follow: lower eyelid scutellation and pigmentation; lateral body folds and lateral granules; dorsal cephalic scutel- lation (prefrontals and supraoculars); femoral pores; body, limb, and lateral cephalic scutellation (labials, tempo- rals ) . On the basis of the first and second of Wagner's criteria, the following char- acter states are judged primitive: pig- mented lower eyelid composed of sev- eral scales; presence of lateral body folds, lateral granules, prefrontal scales, three supraoculars, and femoral pores. Popu- lations of FhoUdohohis with low num- bers of body scales are usually found in cool-wet montane habitats which are recently formed environments. Thus I consider high mean values for body, limb, and lateral cephalic scutellation to be primiti\'e states and reduction of scutellation to be the derived condition. A composite hypothetical ancestral OTU of FhoJidohoJus therefore has the follow- ing character states: three supraoculars, two prefrontals; lateral body folds pres- ent; lateral granules present; femoral pores present; pigmented lower eyelid with several scales; relatively high mean values for body, limb and lateral cephalic scutellation. A comparison of the com- posite ancestral OTU with each of the species of FhoJidoholus reveals that P. affmis is most primitive in its suite of character states (Table 3). FhoJidoho- lus prefrontahs, although exhibiting pre- frontal scales, is relatively derived in showing moderate reduction of body and limb scutellation, having two supraocu- lars, and totally lacking femoral pores. Fhohdohohis monthim exhibits mostly derived states — two supraoculars, total absence of prefrontals and femoral pores. and reduction in lateral granules, body and limb scutellation. FlioUdohoJus mac- ])njdei shows the primitive condition in prefrontal scales and femoral pores among some populations, but this species is relatively more derived than ancestral in having two supraoculars, low mean values in body, limb and lateral cephalic scutellation, and reduction or absence of lateral granules. Fhohdohohis annectens is the most derived and specialized mem- ber of the genus, having two supraocu- lars, a single, usually ti-ansparent lower eyelid disc, and lacking prefrontals, lat- eral body folds and lateral gi-anules. The body and limb scutellation also show considerable reduction. Ecological data support the above arrangement. Fhohdohohis monthim and F. machnjdei exhibit relatively narrow moisture tolerance limits and occur pri- marily in geologically recent habitats — montane forests and wet paramo which developed subsequent to the Andean orogeny. Fhohdohohis prefronfalis oc- curs in relatively dry to xeric regions which are geologically recent, and is con- sidered ecologically specialized. FhoU- dohohis annectens, although morphologi- cally derived and specialized, is associ- ated with some Amazonian floral and faunal components. However, ecological specialization is suggested by the narrow vertical distributional limits; the species does not extend below 2000 m or much above 2335 m. Fhohdohohis affinis, by contrast, is ecologically generalized (shows a broad moisture tolerance), oc- curring in dry, interior basins as well as in mesic areas in the Eastern Cordillera (Bafios, Cerro Llanganate), and de- scending to at least 1800 m. Thus, an affinisAike ancestor would have the greatest probability of having occupied the primitive, pre-Andean lowlands. Although my approach to the esti- mation of primitive character states is subject to criticism by some evolutionary biologists, I believe my position is not only justified but preferred. The associa- tion of a particular set of characters as an integrated unit at the present time 26 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY does not mean that this set coevolved throughout all of history. The probabil- ity of this is very low for it is the change in character complexes and character state combinations that produces various levels of evolutionary diversification. Character states in a variety of combi- nations are obvious among groups of species and higher taxa thus providing ample evidence for potential independ- ence of characters and /or character states at different points in evolutionary time. Thus considering characters po- tentially independent and treating them independently for estimation of ancestral states constitutes a minimum assumption. In two Prim Networks generated from weighted and unweighted data sets of FhoUdoholus, an OTU of P. montiiim represents the cladistic and patristic cen- ter in each case. For reasons presented above, I consider P. 7nonthim to be a relatively derived species. If my inter- pretation is correct, the position of the montium OTU on the Prim Network would indicate evolutionary success, i.e. the species machrydei, prefronfalis, and annectens arose from a montitim-Mke progenitor. The close relationship among these species is apparent because they are linked together on the basis of two supraoculars, absence of prefrontals ( ex- cept prefronfalis), and generally low values for 14 meristic characters of the body and limbs (see Table 3 and Ap- pendix A). On the other hand, ajfinis is distinct in having 3 supraoculars, 2 prefrontals, and high mean values for the meristic characters of the body and limbs. The position of a potential an- cestral OTU relative to the cladistic center of a Prim Network also can be a function of the relative number of re- lated and unrelated OTUs in the analy- sis. The cladistic position would become a sampling phenomenon when using local populations as input OTUs (i.e., including many local populations that are closely related to the potential an- cestor would effect its central position), or would become an evolutionary phe- nomenon when using species or higher categories as input OTUs. Phyletic Constructs and Cliaracter Consistencies. — A total of four Wagner Diagrams \\^as generated from various modifications (see below) of the weighted data set of FhoUdoholus. Ori- entation of the phyletic constructs was achieved by rooting with the composite ancestral OTU. In the first dendrogram generated from a limited data set, a sample of P. machrydei (with prefrontal scales) clustered with P. prefronfalis. The misclustering was corrected by add- ing a morphometric character (HW). An additional problem was encountered with the incorporation of new samples. The sample of P. prefronfalis from near Rio Caiiar fell between the clusters of P. montium and P. affinis. The sample from Rio Caiiar exhibits high mean val- ues for several meristic characters (see Appendix A ) ; apparently the sum of the low weights of the characters overrode the influence of the few, high-weighted diagnostic characters. Correlation anal- ysis revealed that characters ST3 and SF3 are highly correlated with each other and with other limb characters and thus were deleted. The character SBO also was deleted simply because of the lack of significant mean differ- ences among samples. In addition, sig- nificant correlation was noted between SGV and DEL among the majority of samples. Proper clustering of the Rio Caiiar OTU was achieved by deleting either SGV or DEL. The dendrogram without SGV shows a higher consistency value than does the construct lacking DEL. Moreover, the relative weight and unit character consistency of DEL is greater than that of SGV. Therefore, the Wagner Diagram \\\\\\ DEL (lacking SGV) was chosen as most parsimonious (Fig. 8). The total length of the con- struct is 4(S4.7, and its consistency index is 0.488 (a consistency \'alue of LOO would indicate no homoplasy). One can argue that a consistency in- dex of 0.488 demonstrates that there is so much homoplasy present among the MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 27 Table 4. The patristic distance between the closest interspecific denies (A), and the great- est patristic distance between phj'lctically ad- jacent conspecific denies ( B ) . A. P. affitns — P. monihim (Banos — San Antonio) 31.7 F. niontium — P. annectens ( Otavalo— Loja Valley) 42.7 P. niontium — P. prcfrontalis ( Mulalo— Guaranda ) 20.9 P. montium — P. machn/dci (Otavalo— S Cutchil) 55.6 P. prcfrontalis — P. macbnjdei (Guaranda— S Cutchil) 75.4 P. annectens — P. macbnjdei (Loja Valley— E Loja) 85.7 B. P. affinis (SSE Patate— Tixan) 28.3 P. 77}oniiinn (W San Antonio — Cayambe) — _ 17.9 P. prcfrontalis (Guaranda — Cutchil) 31.6 P. machnjdei (ELoja— S Cutchil) 36.2 characters that the phylogenetic conclu- sions drawn are \'ery questionable. I attribute the low consistency index to the use of local population samples as input OTUs. One of the characteristics of local biological populations is their capacity to differ from one another sta- tistically. Any character, especially a meristic one, can be expected to change in mean value from one local sample to the next. Thus, on a phyletic construct, numerous character state reversals are not only possible, but likely, and when many meristic characters are taken to- gether ( as in this data set ) the effect can be compounded, resulting in a relatively low consistency index for the Wagner tree. When all the local population val- ules are averaged into a mean for each species, the number of possible charac- ter steps can be greatly reduced and with little or no effect on the observed range for the character. For example, the unit character consistency for DEL is 0.403 (Table 5) when the input OTUs are local samples. After pooling the sam- ples to obtain a mean for each species, the consistency index for DEL was re- calculated using essentially the same se- quence of OTUs as in the original character state tree, but adjusting the observed range of DEL to correspond to the new mean values. The consistency index thus obtained for DEL was 0.921, indicating little homoplasy! In like man- ner (using species as input OTUs) the overall consistency index for the phyletic construct could be greatly increased. However, I am interested in the relative positions of the local populations from the standpoint of morphological simi- larity. I am not purporting to show evo- lutionary relationships between local populations, but only between local pop- ulation clusters (species); the sequence of the clusters is significant from the standpoint of evolution. Hence, the aforementioned criticism regarding low consistency has no validity here. I con- sider the arrangement in figure 8 to be a reliable estimate of the evolutionary relationships between the species of Fholidohohis. The local samples of FlioJidoholiis on the phyletic construct cluster according to the species recognized herein by the criteria of character concordance and svmpatry. In most cases, the interspe- cific patristic distance is greater than the distance between intraspecific denies (Table 4). A notable exception, how- ever, occurs in PhoJidohoJus prefwntolis. For example, the patristic distances be- tween OTUs from Guaranda (Pacific slope) and Cutchil or NE Cuenca (Cuenca Basin) are greater than the minimum distance between P. montium and P. prefrontaJis. There is no doubt that the samples from the Pacific slopes and the Cuenca Basin are conspecific. Together they form a monophyletic clus- ter, have in common conservative char- acters of head scutellation, and share other characters (keeled dorsals, color pattern), which were not used in the evolutionary analysis. The great intra- specific divergence is attributed to the cumulative difference of several low- weighted, meristic characters. Table 5 shows the relative weight and consistency of each character. Using a nonparametric test (Spearman's co- 28 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Rio Canar annectens Loja Valley E Lnja SW Cajabamba 14 km N Biblidn 9 km N Biblidn prefrontalis montium aff/nis SSE Rotate Ambato SW San Miguelito Chambo Grande S Los Andes iMiini 0 10 20 PATRISTIC DISTANCE Ancestor Fig. 8. A Wagner Diagram based on 29 OTUs (local samples) of PJiolidoboJits. Relationships are drawn to scale. For discussion and interpretation see text and tables 4 and 5. efficient of rank correlation ) , I found no association between the a priori estima- tion of relative weight and unit charac- ter consistency. Thus there is low pre- dictive power between the two param- eters in this data set. Although charac- ters of head scutellation (NSO, NPF) and body-tail scutellation (DEL, SAB, SAT) are more conservative than the others, considerable homoplasy is associ- ated with the majority of characters. As explained earlier, the low character con- sistencies can be attributed to the meris- tic nature of the characters themselves and the use of local biological popula- tions as the input OTUs. The character LEL (a binary state character) shows no homoplasy. On a probability basis, MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 29 one would expect different saurian groups to share the same or similar average numbers of scales more fre- quently than to share a structural speciali- zation. The high weight but low^ con- sistency of NPF suggests that it is adap- tive (or at least associated with an adaptive trait), and has been subjected to changing selection pressures relatively frequenth' through e^•ok^tionary time. The nature of prefrontal scales should not be used as a generic character in the taxonomv of microteiids, because the scales may be non-homologous even among individuals within a local sample. This conclusion follows from differences noted in the position of the scales rela- tive to the anterior corner of the ocular orbit. In some specimens the scales have been formed by fragmentation of the frontal plate, but in other specimens the scales have been derived largely or entirely from the supranasal plate. For- mation of the prefrontal scales has not involved changes in configuration of cranial elements. Regression of head length (ear to snout terminus) against snout length (eye to snout terminus) in samples with and without prefrontal scales resulted in a tight cluster of points with no significant differences in slope or y-intercept. Comparisons have been based on samples of P. machrydei with and without prefrontal scales, P. prefron- talis and P. monthim, and P. affinis and P. moniium. Evoliifionanj Perspective. — The rela- tive positions of the species clusters on the Wagner Diagram (Fig. 8) indicate that an a;^n/.s-]ike ancestor gave rise to a derived, generalized monfitim-\ike stock; the latter, in turn, underwent evolutionary radiation producing several ecologically or morphologically special- ized species. Essentially the same cla- distic result is obtained from the four dendrograms generated; they differ only in the relative positions of local samples within a given species cluster. E\'olu- tionary interpretation is most meaningful at the level of species clusters. I do not attach evolutionary significance to the Table 5. The relative weight and consistency of each character used in the Quantitative Phy- letic Analysis of Pholidobolus. Character Weight Consistency NPF 3.39 0.304 NSO 2.69 0.567 SUJ 1.3S 0.309 SLT 1.18 0.352 SGI 1.83 0.318 DEL 2.42 0.403 NTS 1.31 0.286 SAB 2.46 0.411 SAT 2.29 0.470 SAF 1.94 0.398 SF5 0.96 0.408 ST4 1.50 0.412 ST5 1.23 0.340 FP 1.64 0.476 LG 0.53 0.357 LEL 0.65 1.000 LES 1.05 0.457 HW 1.18 0.780 sequence of local samples within a clus- ter. Earlier discussion revealed that mor- phological closeness of geographically distant denies is apparently a reflection of climatic similarities between localities. Therefore, patristic relationships be- tween intraspecific samples are best re- garded simply as measures of morpho- logical affinity. Probably all high northern Andean biotic patterns have developed since the end of the Tertiary, and most have been explained in terms of Pleistocene cli- matic events (Haffer, 1970; Vuilleumier, 1971). The great influence of glacial- interglacial oscillations on plant and animal distributions has been the funda- mental assumption underlying the inter- pretation of speciation patterns. During glacial periods, areas under permanent ice are thought to have increased. Pre- sumably this resulted in the lowering of vegetati\^e zones, and thereby permitted high montane organisms to disperse through suitable habitat to adjacent mountain ranges. During interglacial periods, intermontane valleys and passes became arid; thus, xeric-adapted forms were allowed to spread, whereas mesic- adapted, montane organisms were re- stricted or isolated. Admittedlv, this in- 30 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY terpretatioii is simplistic. It does not take into consideration such factors as physiogi^aphic features influencing pre- cipitation and drainage patterns, or minor climatic oscillations which must ha\'e occurred during the major glacial and inter glacial periods. Therefore, the picture of climatic change, dispersal, and evolution of Andean organisms during the Pleistocene probably is more compli- cated than generally has been thought. There is evidence that the distribu- tional patterns of PhoUdohohis were modified by climatic oscillations during the Pleistocene. The southern isolate of PhoUdohohis affmis near Saraguro indi- cates that the distribution of this species was more widespread than it is at pres- ent. Perhaps during a dry climatic pe- riod, populations of the derived, xeric- adapted Phohdoholus prefwntalis dis- persed southward along the Pacific slopes and through the Cuenca Basin, displac- ing the ecologically generalized P. affinis from most of its former range. This ex- planation is considered the simplest in- terpretation of the present distributional relationships of the two species. During one or more wet climatic periods, mesic- adapted, monthim-Mke populations (pro- genitors of P. machnjdei) were able to spread southward along the Eastern and Western Cordilleras. During dry periods, these populations could have survived in mesic, high slope and crest situations, especially along the Amazonian slopes of the Eastern Cordillera. PJwhdohohis annectens, a species endemic to the southern Loja Basin, is a dispersal prod- uct of the montium-MVe progenitor popu- lations. Macroplwhdus nitlweni, appar- ently a highly specialized derivative of PhoUdohohis stock, provides evidence that the southward dispersing progeni- tors reached northern Peru. Vuilleumier ( 1971 ) placed great em- phasis on isolation by geographical and ecological (glacial and vegetative) bar- riers as the mechanism for speciation in high Andean organisms during the Pleis- tocene. Major evolutionary diversifica- tion of PJwUdoI)ohis probably occurred during this time. However, I do not consider isolation necessary in explaining speciation in PhoUdohohis. Ehrlich and Raven (1969) provided evidence sup- porting their contention that natural se- lection is the major e\'olutionary force and that ultimatelv, it determines what effect gene flow will have between popu- lations. In PhoUdoholus several lines of evidence support this argument. First, there is no correlation (rrir 0.064) be- tween geographical distance and mor- phological similarity among inti-aspecific local samples. If gene flow were main- taining the similarity between popula- tions, one would expect adjacent sam- ples to be more similar to each other than to progressively distant samples. Second, maximal glacial extension de- picted by Saucr (1965) would not have formed complete barriers betvveen the interandean basins occupied by PhoU- dohohis. Montane passes apparently existed between the Quito and Lata- cunga basins, the Riobamba and Alausi basins, and between the Cuenca and Alausi basins. Third, the existence of vegetative barriers between incipient species of PlwUdoholiis is unlikely. The moisture tolerance limits of the species overlap in part and each species (except P. annectens) inhabits several vegetative associations. There is little reason to believe, for example, that the evolution of P. machnjdei and P. prefrontaUs from a common stock inxolved ecological iso- lation. Ancestral populations in the dry Alausi and Cuenca basins could have diverged from populations on the adja- cent, wet slopes in response to diff^eren- tial selection pressures while maintaining parapatry (parapatric speciation). The probable effect of glacial-interglacial os- cillations simply would have been to displace vertically the zone of contact. The divergence of P. monfiinn from an «j^;i/.s-like progenitor can be argued simi- larly. Therefore, speciation in PhoU- dohohis can be best explained in terms of different selective regimes acting upon the populations through time. MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 31 ACCOUNTS OF SPECIES The genus Fholidohohis is known only from the Eeuadorian Andes. De- tailed locality data for each species are presented in appendix B. Distribution maps are shown in figures 6 and 7. Pholidobolus Peters, 1862 Pholidoholus Peters, 1862, Abh. Akad. Wiss. Berlin, 1862:195 (Type species.— £c- pleopus { Pholidobolus ) montium Peters, 1862, by monotypy ) . AspidoJacmus Peters, 1862, Abh. Akad. Wiss. Berlin, 1862:199 (Type species. — Ec- pleopus (Aspidolaemus) affinis Peters, 1862, by monotypy). Diagnostic Definition. — 1) Teiid liz- ards of Group II (Boulenger, 1885) hav- ing snout-vent lengths of 66 mm or less; 2) tail accounting for 69 percent or less of total length; 3) limbs pentadactyl, digits clawed; 4) body and tail cylindri- cal, neck not greatly constricted; 5 ) head distinctly flattened, depth 59-73 (67.7) percent of width; 6) tympanum deeply recessed; 7) snout obtusely pointed; 8) nostril piercing nasal suture; 9) dorsal scales imbricate and weakly keeled to striated with some placoid; 10) dorsal scales in transverse series; 11) dorsal scales longer than wide, quadrangular to subhexagonal; 12) gular scales smooth, imbricate, becoming larger posteriorly; 13) two medial rows of widened gulars anterior to collar fold; 14) ventral scales smooth, imbricate; 15) ventral scales rec- tangular, in transverse and longitudinal series; 16) tongue covered with imbri- cate, scale-like papillae; 17) hemipenes with minute calcareous spines; 18) 26-27 presacral vertebrae; 19) 21-23 ribs; 20) 5-6 pairs of ribs attached to pectoral girdle; 21) one pair of abdominal ribs attached to each other midventrally, pos- terior to the xiphisternum. KEY TO THE SPECIES OF PHOLIDOBOLUS 1. Three supraoculars, middle one wedged between first and third with apex usually reaching superciliary row (Fig. 5A); two prefrontals P. affinis Two supraoculars, subequal in size 2 2. Prefrontals present 3 Prefrontals absent 4 3. Dorsolateral pale stripe distinct, ex- tending to tip of snout; femoral pores absent in both sexes; head not dis- tinctly broader in males than in fe- males; sides of neck and tail without red stripe ( orange-brown in preserva- tive) in males P. prefrontalis Dorsolateral pale stripe distinct, not extending to tip of snout; femoral pores usually present in males; head distinctly broader in males than in females; sides of neck and tail with red stripe in males P. macbnjdei 4. Lower eyelid usually having a single transparent disc; lateral granules usu- ally absent at midbody; dorsum uni- formly gray-brown P. annectens Lower eyelid having 3-6 opaque scales; lateral granules usually pres- ent at midbody 5 5. Dorsolateral pale stripe distinct, not extending to tip of snout; femoral pores usually present in males; head distinctly broader in males than in females; sides of neck and tail with red stripe in males P. macbnjdei Dorsolateral pale stripe distinct, ex- tending to tip of snout; femoral pores absent in both sexes; head not dis- tinctly broader in males than in fe- males; sides of neck and tail without red stripe in males P. montium Pholidobolus affinis (Peters) New combination Ecpleopus {Aspidolaemus) affinis Peters, 1862, Abh. Akad. Wiss. BerHn, 1862:199, pi. 3, fig. 1 [Holotype.— ZSM 664/0 from Pichincha Prov., Ecuador].^ ^ Through the courtesy of Thomas Uzzell, I recently examined the type specimen (ZMB 16593) oi Prionodactyhis ocelUfer Werner, 1901. Uzzell and I agree that Prionodactylus ocellifer is a junior synonym of Ecpleopus {Aspidolae- mus) affinis Peters, 1862. 32 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Ecpleopus affinis — Boulenger, 1885, Cat. Liz. Brit. Mus., 2:402. Aspidolaemus affinis — Uzzell, 1969, Postilla, 135:21. Diagnosis. — 1) Head \\'idth not sex- ually dimorphic; 2) head width/ snout- vent length ratio 0.151-O.lSO (0.162); 3) three supraoculars, the middle one being wedged between first and third with apex usually reaching superciliary row; 4) usually two prefrontals; 5) femoral pores present in about two percent of males; 6) lower eyelid scales opaque, three to six in number; 7) dorsals keeled to striated, seldom smooth; 8) lateral granules usually present at midbody; 9) lateral body fold present; 10) dorsolat- eral pale stripe extending to tip of snout; 11) body having black reticulations lat- erally on reddish orange ground color, lateral ocelli usually present; 12) sides of neck and tail without red stripe; 13) ven- ter yellow in males; 14) venter pale yel- low to gray in females; 15) underside of tail orange to orange-red in males. Description and Color Pattern (140 in- divichiahj. — Maximum snout-vent length 64 mm for males, 58 mm for females. For details of scutellation and proportions see table 3 and generic definition. Dorsum gray-brown to olive-brown with or without a dark brown to black middorsal stripe, or traces of it; dorso- lateral stripe distinct, \\'hite to cream or yellow fading to pale gray posteriorly, edged with black, extending from snout along edge of orbit, terminating about midbody, in some individuals extending to near base of tail; labial stripe pale yellow or cream extending along upper lip to shoulder; side of head between dorsolateral and labial stripes uniform pale gray to dark brown; gular region dull white to yellowish; limbs gray- brown with fine black reticulation. Males: Venter yellow to orange-red with scattered dark flecks posteriorly in some indixiduals; underside of tail orange to orange-red with or without brown or black flecks; flanks pale orange to orange-red with black reticulation re- sulting in indistinct ocelli; ventrolateral orange-red wash intensified in breeding males, extending high on flanks and dor- sal surface of tail; iris tan to pale gold (PI. lA). Females: \^enter and underside of tail uniform pale yellow to pale or dark gray; usually orange-red color lacking; flanks pale brown, reticulated with black, or uniform dark brown or black; dorsal surface of tail brown to pale yel- low. Color differences among some popu- lations are evident. Indi\'iduals from near Tixan have an orange-red, instead of yellow, venter; yellow is present on the chest of some individuals. Lizards from near Banos are noticeably pale; the venter is very pale yellow, and males have only faint traces of orange on the flanks. The nature of the middorsal stripe varies in local populations. For example, in 38 indi\'iduals from Mulalo, the stripe is distinct in 34, faint in three, and absent in one; in 16 indixiduals from southeast of Patate, the stripe is distinct in five, faint in nine, and absent in two individuals. One specimen from Saraguro differs from all others examined. The belly and subcaudal areas are gray; the dorsum is tan, and the side of the head and neck are dark brown. Distribution. — PlioIidohoJtis affinis oc- curs principally in the Latacunga and Riobamba basins of the Central Valley. Mulalo is the northernmost record for the species. A single specimen taken at Saraguro probably represents a disjunct, relictual population. Plwlidoholus af- finis reaches the Pacific slopes of the Western Cordillera, penetrating at least as far as 3.3 km south of Tixan. Pholidobolus prefrontalis new species Hohtype.—KU 141093 from 4.9 km, by road, south of Tixan, Chimborazo Province, Ecuador; obtained by R. R. Montanucci on 3 June 1971. Parati/pes.—KV 141094-141113, CAS 132583-84, USNM 193219-220 from 3.3- 4.9 km south of Tixan, Chimborazo Province, Ecuador. MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 33 Diagnosis. — 1) Head width not sex- ually dimorphic; 2) head \\idth/ snout- vent length ratio 0.150-0.177 (0.164); 3) two supraoculars, subequal in size; 4) usually two prefrontals; 5) femoral pores absent in both sexes; 6) lower eyelid scales opaque, three to six in number; 7) dorsals usually keeled; 8) lateral granules usually present at midbody; 9) lateral body fold present; 10) dorso- lateral pale stripe extending to tip of snout; 11) sides of body with single, broad brown stripe, bordered by a pale stripe above and below, or several alter- nating brown and cream longitudinal stripes; 12) sides of neck and tail with- out red stripe; 13) venter gray to blue- black posteriorly in males; 14) venter gray or salmon in females; 15) underside of tail blue-black in males. Description of HoJotype. — Male, snout-vent length 50 mm; tail length 75 mm, partly regenerated; two prefrontals; four scales between the orbits; two su- praoculars; 12 scales along upper lip; 12 scales along lower lip; 17 scales from mental to collar fold; 28 venti-als; 39 dorsals; six temporals; 39 scales around bodv; 19 scales around tail; 23 scales along forelimb; eight scales on third finger; six scales on fifth finger; 10 scales on third toe; 13 scales on fourth toe; nine scales on fifth toe; femoral pores absent; lateral granules present; lower eyelid opaque; four lower eyelid scales. Dorsum uniform pale gray-brown; dorsolateral stripe pale yellow, extending from tip of snout to above forelimb; la- bial stripe white, extending to shoulder; sides of head and neck dark l^ro^^'n; flanks with single broad, brown stripe; narrow pale line extending from inser- tion of forehmb to hind limb; sides of tail with broad, brown stripe; fight brown stripe extending dorsally on tail; chin, throat and chest salmon, blending into blue-black on belly and under tail (PLIC). Description and Color Pattern (109 in- dividuals).— Maximum snout-vent length 57 mm for males, 63 mm for females. For details of scutellation and propor- tions see table 3 and generic definition. Dorsum pale gray to dark brown ( coppery cast evident in most young and sub-adults), with or without a dark brown to black middorsal stripe, or traces of it; middorsal brown zone with or without yellowish flecks; dorsolateral stripe white to yellow, edged with black dorsally, terminating above shoulder or grading imperceptibly into ground color near midbody; labial stripe yellow to white, terminating at forearm; pale cream lateral stripe extending beyond forearm to hind limb; single broad brown to dark brown stripe between dorsolateral and lateral cream stripes, or several brown stripes alternating with cream stripes; belly pink to salmon, or gray to blue-black with or without dark spots or flecks laterally and posteriorly; chin white to gray; underside of tail gray to blue-black. Considerable variation exists in belly coloration between local populations and can be associated with geographic area. Among the populations on the Pa- cific slope, the belly color varies from gray-white (Guaranda) to fight gray with pinkish cast ( San Jose del Chimbo, two specimens ) to a rich salmon ( Alausi and Tixan). Males from the Alausi and Tixan populations have considerable blue-black or gray on the posterior belly, with salmon being restricted to the chest and throat. The belly color of the popu- lation at Caiiar is pale gray. Varying amounts of venti'al spotting are present among the samples from the Pacific slopes. At Guaranda, 26 individuals lack spots; seven have spots posteriorly. Most individuals in populations from the Cuenca Basin lack spotting, the venter being yellowish to dull white. Polymorphism occurs in the nature of the middorsal stripe and number of lateral brown stripes (Table 6). In gen- eral, the samples from the Cuenca Valley have more brown stripes than those from the Pacific slopes. The amount of strip- ing in a local sample apparently is asso- ciated with moisture. Prominently striped indi\ iduals are usually found at 34 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY mesic localities characterized by dense vegetation. More uniformly colored liz- ards occur in xeric, exposed areas. The color pattern probably has a cryptic function responding directly to preda- tion. The pattern may also be in\'olved with thermoregulation. Distribution. — PJioIidobolus prefron- talis occurs along the Pacific slopes of the Western Cordillera from Guaranda south to Canar; populations are also found throughout the Cuenca Valley. Etymology. — The specific name re- fers to the prominent prefrontal scales present in this species. Pholidobolus montium (Peters) Ecpleopus {Pholidohohis) ryiontiiim Peters, 1862, Abh. Akad. Wiss. Berlin, 1862: 196, pi. 2, fig. 3. Pholidobolus montium — Boulenger, 1885, Cat. Liz. Brit. Mus., 2:403. Types. — The two syntypes used in the description of PJioIidobolus montium represent two species. RMNH 3401 is hereby designated as the lectotype, be- cause this specimen is from a known locality (Quito) and because Peters (1862) illustrated this specimen and used it for most of the type description. The other syntype (ZMB 900) from "western Ecuador" belongs to the species described in the next account. Diogno.sis. — 1) Head width not sex- ually dimorphic; 2) head width/ snout- vent length ratio 0.155-0.175 (0.163); 3) two supraoculars, subequal in size; 4) prefrontals absent; 5) femoral pores ab- sent in both sexes; 6) lower eyelid scales opaque, three to six in number; 7) dor- sals striated to smooth; 8) lateral gran- ules present or absent at midbody; 9) lateral body fold present; 10) dorsolat- eral pale stripe extending to tip of snout; 11) several brown and cream longitudi- nal stripes on sides of body; 12) sides of neck and tail without red stripe; 13) venter light gray in males; 14) venter pale yellowish white to pale gray in females; 15) underside of tail gray to blue-black in males. Description and Color Pattern (138 in- dividuals).— Maximum snout-vent length 56 mm for males, 66 mm for females. For details of scutellation and propor- tions see table 3 and generic definition. Dorsum pale gray to dark brown, with an iridescent coppery cast in many indi^'iduals; vertebral area with or with- out brown to black middorsal stripe, or brown to black flecks; dorsolateral stripe pale yellow, edged with dark brown or black, extending to above shoulder, or fading into dorsal dark field near mid- body; labial stripe cream to white, ex- tending to forearm; sides of head be- tween labial and dorsolateral stripes uni- form dark brown to black; sides of body with one or several brown to black- brown stripes alternating with several Table 6. Polymorphism in lateral and middorsal stripes in samples of PJioJidohohis prefronialis. Frequency of each morph is expressed as a percentage of the sample size at each locality. Sample size is indicated in parentheses. Number of dark lateral stripes Sample 12 3 4 Guaranda (33) 21.2 Alausi (16) 100 Tixan (24) 100 Cutcliil (13) Nature of middorsal stripe Absent Trace Present 57.6 21.2 57.6 36.3 6.1 50.0 50.0 70.9 20.8 8.3 7.8 61.5 30.7 7.7 23.1 69.2 Plate 1. A. Pholidobolus affinis $ (KU 140983; 55 mm S\'L) from 8.9 km SSE Patate; B. P. mon- tium S (KU 141081; 55 mm SVL) from 2.5 km E Rio Chiche; C. P. prefiontalis 6 ( KU 141093, holotype; 50 mm SVL) from 4.9 km S Tixan; D. P. machnjdei $ (KU 140993, holotype; 53 mm SVL) from 9.1 km N, 2.3 km W Biblian; E. P. annectens 9 (KU 140986; 55 mm S\'L) from Loja Valley. Photos liy George R. Pisani. PLATE 1 MONTANUCCI: ANDEAN LIZARD GENUS PHOLIDOBOLUS 35 T.\BLE 7. Polymoiphism in lateral stripes in samples of Pholidobolus montium. Frequency of each moiph is expressed as a percentage of the sample size at each locality. Sample size is indicated in parentheses. Sample Ibarra (20) Otavalo (33) Cayambe (38) W San Antonio (20) San Antonio (17) Quito (14) Lago Cuicocha (20) E Rio Cliiche (31) . Number of dark lateral stripes 2 3 4 5 10.0 35.0 28.9 28.9 5.0 5.0 17.6 28.5 6.4 9.7 50.0 5.0 63.6 36.4 34.2 8.0 60.0 30.0 64.8 17.6 71.5 5.0 10.0 12.9 61.3 80.0 9.7 5.0 white, cream or yellowish stripes (Pi. IB); chin and throat dull white to pale gray; venter pale yellowish white to iridescent pale gray, with or without black spots or flecks posteriorly and lat- erally; underside of tail gray to blue- black. The number of lateral brown stripes is variable; frequencies of various num- bers in different local populations are shown in table 7. The number of lateral stripes may be associated with moisture and vegetative characteristics of the lo- cality, presumably in response to preda- tor pressure. However, the relationship between amount of striping and ^'egeta- tion is not as ob\'ious as in Fholidoholus prefrontal is. Polymorphism in the nature of the middorsal .stripe and ventral pat- tern is summarized in table 8. There is no evidence to indicate that venti'al spot- ting ^'arics ontogenetically. The insular population from Lago Cuicocha shows slight differences in ventral coloration compared to surrounding samples. The chest color is pale orangish yellow in five of eight males examined. Among 12 females, only three have the yellowish color, whereas nine have the t\'pical grayish color with or without an irides- cent cast. DhiriJmfion. — PlwJidohohis montium occurs in the northern basins of Quito, Ibarra, and adjacent valleys, and is ex- pected in extreme southern Colombia. The southern limit of its disti^ibution is near the northern end of the Central Valley (Latacunga Basin) near Mulalo. A more southern record from near Bafios needs confirmation; I searched the Bafios area unsuccessfully for additional speci- mens. Pholidobolus macbrydei new species HoIofype.—KV 140993 from 9.1 km north and 2.3 km west (by road) of Biblian, near Hacienda Oeste, 3070 m, Table 8. Pohmorphism in middorsal stripe and ventral pattern in samples of Fholidoholus montium. Frequency of each morph is expressed as a percentage of the sample size at each locality. Sample size is indicated in parentheses. Sample Nature of Absent middoi Trace rsal stripe Present Uniform light gray Ventral pattern Posterior Totally half spotted spotted Uniform black Ibarra (20) 20.0 60.0 54.5 31.6 55.0 58.8 35.7 30.0 87.1 20.0 45.5 28.9 35.0 11.8 50.1 65.0 3.2 60.0 36.4 76.3 45.0 94.1 21.4 75.0 19.3 25.0 48.5 23.7 50.0 5.9 42.9 25.0 71.0 15.0 9.0 5.0 35.7 97 Otavalo (33) 6.1 Cavambe (38) 39.5 W San Antonio (20) ... San Antonio ( 17) ... 10.0 29.4 Quito (14) 14.2 Lago Cuicocha (20) ... E Rio Chiche (31) ..... ... 5.0 ... 9.7 36 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Canar Province, Ecuador; obtained by R. R. Montaniicci and B. MacBrvde on 23 May, 1971. Paratypes.—KU 140994-141014, CAS 132581, USNM 193217 from 14.2 km north (by road) of Biblian, 3430 m, and KU 141015-141027, CAS 132582, USNM 193218 from 9.1 km north and 2.3 km west (by road) of Biblian, near Ha- cienda Oeste, 3070 m, Cafiar Province, Ecuador. Dia