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HARVARD UNIVERSITY

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Museum of

Comparative Zoology

The Great Basin Naturalist

VOLUME 35, 1975

Editor: STi:i'Hr.N L. W(

Published at Brigham Young University, by Brigham Young University

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TABLE OF CONTENl S

Volume 36

Number 1 - March 31. 1975

Evolution of the sceloporine lizards (Iguaniclae). Kenneth R. Larsen

and Wilmer W. Tanner 1

New synonymy and new species of American bark beetles (Coleop-

tera: Scolytidae) . Stephen L. Wood 21

Genetics, environment, and subspecies differences: the case of Polites

sabuleti (Lepidoptera: Hesperiidae). Arthur M. Shapiro 33

Life history and ecology of Megarcys signata (Plecoptera: Perlodidae),

Mill Creek, Wasatch Mountains, Utah. Mary R. Gather and Arden

R. Gaufin 39

Records of stoneflies (Plecoptera) from Nevada. Marv R. Gather, Bill

P. Stark, Arden R. Gaufin ' 49

Growth of Plecoptera (stonefly) nymphs at constant, abnormally high

temperatures. Joseph M. Branham, Arden R. Gaufin, and Robbin

L. Traver 51

Water balance and fluid consumption in the southern grasshopper m.ouse,

Onychomys torridus. Vernon C. Bleich and Orlando A. Schwartz ... 62 A systematic study of Coenia and Paracoenia (Diptera: Ephvdridae).

Wayne N. Mathis 1 65

Environmental factors in relation to the salt content of Salicornio pa-

cifica var. utahensis. D. J. Hansen and D. J. Weber 86

New records of stoneflies (Plecoptera) from New Mexico. Bill P. Stark,

Theodore A. Wolff, and Arden R. Gaufin 97

The authorship and date of publication of Siren intermedia (Amphibia:

Caudata). Hobart M. Smith, Rozella B. Smith, and H. Lewds

Sawin ..-. 100

New mites from the Yampa Valley (Acarina: Cryptostigmata: Ori-

batulidae, Passalizetidae). Harold G. Higgins and Tvler A.

Woolley 103

The identity of Boucourt's lizard Eunieces capito 1879. Hobart M. Smith,

Rozella B. Smith, and Jean Guibe 109

Studies in nearctic desert sand dune Orthoptera. Part XV. Eremog-

raphy of Spaniacris with biological notes. Ernest R. Tinkham 113

Roosting behavior of male Euderma maculatum from Utah. Richard

M. Poche and George A. Ruffner 121

The nest and larva of Diploplectron hrunneipes (Cresson) Hymenop-

tera: Sphecidae). Howard E. Evans 123

Number 2 - .Tune 30, 1975

A revision of the Phacelia Crenulatae group (Hydrophyllaceae) for

North America. N. Duane Atwood 127

Rodent populations, biomass, and community relationships in Arte- misia tridentata. Rush Vallev, Utah. D. W. Nichols, H. D. Smith, and M. F. Baker ." 191

Computerized reduction of meteorologic measurements from irrigated and nonirrigated plots in central Utah. Ferron L. Andersen and Paul R. Roper - 203

3-^ '

Clarence Cottani, 1899-1974; a distinguished alumnus of Brigham

Young University. Vasco M. Tanner 231

Evolutionary divergence in closely related populations of Mimulus guttatus (Scrophulariaceae). Karen W. Hughes and Robert W. Vickery, Jr 240

Number 3 - September 30, 1975

Urosaurus and its phylogenetic relationship to Uta as determined by osteolog}' and myology (Reptilia: Iguanidae). Charles Fanghella, David F. Avery, and Wilmer W. Tanner 245

Distribution and adundance of the black-billed magpie {Pica pica) in

North America. Carl E. Bock and Larry W. Lepthien 269

Nectar composition of hawkmoth-visited species of Oenothera (Ona-

graceae). Robert E. Stockhouse, II 273

A revision of the nearctic species of Clinohelea Kieffer (Diptera: Cera-

topogonidae). William L. Grogan, Jr., and Willis W. Wirth 275

Basidiomycetes that decay junipers in Arizona. R. L. Gilbertson and

J. P. Lindsey 288

Body size, organ size, and sex ratios in adult and yearling Belding

ground squirrels. Martin L. Morton and Robert J. Parmer 305

Photoperiodic responses of phenologically aberrant populations of pie- rid butterflies (Lepidoptera). Arthur M. Shapiro 310

Additional records of reptiles from Jalisco, Mexico. Philip A. Medica,

Rudolf G. Arndt, and James R. Dixon 317

Invasion of big sagebrush {Artemisia tridentata) by white fir {Abies concolor) on the southeastern slopes of the Warner Mountains, California. Thomas R. Vale 319

Morpholog}' of ephemeral and persistent leaves of three subspecies of big sagebrush grown in a uniform environment. W. T. McDonough, R. 0. Harniss, and R. B. Campbell 325

Number 4 - December 31, 1975

Endangered, threatened, extinct, endemic, and rare or restricted Utah vascular plants. Stanley L. Welsh, N. Duane Atwood, and James L. Reveal ". 327

Utah plant novelties in Cymopterus and Penstemon. Stanley L. Welsh.... 377

The Zygoptera (Odonata) of Utah with notes on their biology. A. B.

Provonsha :. 379

New synonymy and new species of American bark beetles (Coleop-

tera: Scolytidae), Part II. Stephen L. Wood 391

Correlates of burrow location in Beechey ground squirrels. Donald H.

Owings and Mark Borchert 402

Arachnids as ecological indicators. Dorald M. Allred 405

Notes on the genus Bomhylius Linnaeus in Utah, with key and descrip- tions of new species (Diptera: Bombyliidae). D. Elmer Johnson and Lucile Maughan Johnson 407

Breeding range expansion of the starling in Utah. Dwight G. Smith .... 419

Some parasites of paddlefish {Polydon spathula) from the Yellowstone

River, Montana. Lawrence L. Lockard and R. Randall Parsons .... 425

Reproductive cycle of the Belding ground squirrel [Sperrnnphilus hel- dingi) : seasonal and age differences. Martin L. Morton and John S. Gallup 427

A new combination in Penstemon (Scrophulariaceae). Stephen L. Clark.. 434 Some relationships between water fertility and egg production in brown

trout (Salmo trutta) from Montana streams. Lawrence L. Lockard.. 435 Some relationships between internal parasites and brown trout from

Montana streams. Lawrence L. Lockard, R. Randall Parsons, and

Barry M. Schaplow 442

Sexual dimorphism in malpighian tubules of Pteronarcys californica

Newport (Plecoptera) . Ralph R. Hathaway 449

New records of the bat Plecotus phyllotis from Utah. Richard

M. Poche 451

GREAT BASIN NATURALISl

ime35Na1 March 31, 1975

Brigham Young Universit:

MUQ. COMP. ZOOU L.JBRARY

JUN 1 1 1975

HARVARD UNIVERSITY

i ^\

GREAT BASIN NATURALIST

Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo,

Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Zoology; Stanley L.

Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, dean. College of Biological and

Agricultural Sciences; Ernest L. Olson, director, Brigham Young University Press,

University Editor.

The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been continuously published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only original, previously unpublished manuscripts pertaining to the biological natural history of the Great Basin and western North America will be accepted. Manuscripts are subject to the approval of the editor.

Subscriptions. The annual subscription is $9 (outside the United States $10). The price for single numbers is $3 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back numbers should be directed to Brigham Young University Press, Marketing Department, 204 UPB, Provo, Utah 84602.

Scholarly Exchanges. Libraries or other organizations interested in obtaining this journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602.

Manuscripts. All manuscripts and other copy for the Great Basin Naturalist should be addressed to the editor as instructed on the back cover.

The Great Basin Naturalist

Published at Provo, Utah, by Brigham Young University

Volume 35

March 31, 1975

No. 1

EVOLUTION OF THE SCELOPORINE LIZARDS (IGUANIDAE)

Kenneth R. Larsen^ and Wilmer W. Tanner^

Abstract. Phylogenetic relationships among Sceloporine genera are briefly discussed. Species re- lationships witliin the genus Sceloporus are analyzed, and evolutionary lines of descent are proposed.

The genus Sceloporus is composed of three monophyletic groups: Group I, the most primitive, prob- ably developed from Salor-\\ke ancestral stock in Miocene times. This group speciated from stock similar to Sceloporus gadoviae in southern Mexico to S. merriami in the North and contains 7 species in 3 species groups. We propose that these species be included in the genus Lysoptychus Cope. Group II arose from Group I and evolved from centrally located Sceloporus pictus in all directions throughout Mexico. This intennediate group contains approximately 19 species in 5 species groups. Group III also arose from the primitive stock of Group I and radiated from several desert refugia created by Pleistocene glaciation. Evolution of this group in Mexico was generally from north to south with Sceloporus malachiticus extending as far south as Panama. This group contains approximately 33 spe- cies in 5 species groups.

In a previous paper (Larsen and Tan- ner, 1974) we presented our analysis of the species in the lizard genus Sceloporus. Numerical statistical methods were used to analyze the species in the genus Scel- oporus using cranial osteology, external meristic and numeric characters, karyol- ogy, display behavior, and geographic dis- tribution. A new classification for the genus was proposed with three major branches or groups. Group I contained 7 species in 3 species groups. Group II con- tained approximately 19 species in 5 spe- cies groups. Group III contained approxi- mately 33 species in 5 species groups. This classification was supported by the cluster analysis of several different sets of data. Cranial osteology, zoogeograph}', behavior, and karyology were shown to be taxon- omically significant as numeric charac- ters. Stepwise discriminate analysis showed that this classification of the spe- cies of Sceloporus into 3 major groups and 13 species groups was significant at the .999 confidence level.

The purpose of this paper is to present our views on the evolution of the species in the genus Sceloporus. We also propose a ph3dogeny of closely related (Scelop-

orine) genera. We are grateful for the assistance of H. M. Smith, C. C. Carpen- ter, W. P. Hall, and the following per- sons at Brigham Young University: A. L. Allen, F. L. Anderson, J. R. Murphy, M. S. Peterson, J. K. Rigby, N. M. Smith, D. A. White, and S. L. Wood.

Intergeneric Phylogeny

In 1828 Weigmann described several genera, including Sceloporus (S. torqua- tus) . He distinguished Sceloporus from the South American Tropidurus mainly on the basis of femoral pores (S'c^j/o^ thigh, porus=\)OYe) . In 1852 Baird and Girard described the genus Uta (U. stansburiana) which is distinguished from the smaller species of Sceloporus by its gular fold and granular dorsal scales. In 1854 Hallowell erected the genus Urosaurus (U. gracio- sus), which is similar to Uta but has sev- eral rows of enlarged, carinate, imbricate vertebrals or paravertebrals. Two years later Dimieril (1856) described the genus Phymatolepis (Urosaurus bicarinatus) on the basis of enlarged paravertebrals. In 1859 Baird placed Hallowell's genus Uro- saurus in synonymy with Uta, and in

V07 North 500 West, Provo, Utah 84601 .

-Department of Zoology, Brigham Young Universitj-. Provo. Utah 84602.

GREAT BASIN NATURALIST

Vol. 35, No. 1

1864 Cope did the same with Dmneril's Phymatolcpis. Boulenger (1885) raised Cope's Uta thalassina to generic status (Petrosauriis) , but Cope (1900) rejected this proposal and made Petrosaurus a third synon;y^n of Uta. In 1888 Cope erected the genus LrsoptycJius (L. Iateralus^= Sceloporus couchi) on the basis of a single specimen that appeared to have a well- developed gular fold. Subsequent investi- gation (Stejneger, 1904) showed the "gu- lar fold" to be an artifact of preparation on a single specimen which "was pre- served in such a manner as to make a fold across the neck, which formed the basis for the erection of the genus" (Smith, 1939, p. 242). Dickerson (1919) de- scribed the genus Sator (S. grandaevus) which has persisted despite Sator's close similarity to Uta, Urosaurus and Scel- oporus. In 1942 Mittleman resurrected the genera Urosaurus and Petrosaurus. He also erected the genus Streptosaurus based on Uta mearnsi, which is most similar to Petrosawus. He proposed that Uta, Uro- saurus, and Sator all arose independently from Sceloporus. He placed PJirynosoma with the above genera in a distinct group. Smith (1946) moved Sauromalus and Dipsosaurus to more primitive positions but otherwise retained Mittleman's ar- rangement. Savage (1958) placed Strept- osaurus in synonymy with Petrosaurus. He separated Uta from Urosaurus mainly on the basis of sternal and costal mor- phology. He placed Uta and Petrosaurus with the sand lizards (Holhrookia, Unia, and Callisaurus) , leaving Sceloporus, Sa-

SCEIOPORUS

GROUP III

SCELOPORUS

\ GROUP II

COPHOSAURUS HOIBROOKIA

\/

\ /►'^ElOPORUS / GROUP 1

Y

SATOR UMA \

~\

/^^CALIISAURUS

1

^/

/Urosaurus

PHRYNOSOMA^"-\^ "'* \

/

^x

petrosaurus

ANCESTRAL STOCK

Fig. 1. Phylogeny of sceloporine genera and the three major groups in Sceloporus.

tor, and Urosaurus together. Etheridge (1964) rejected Savage's wide separation of Uta and Urosaurus, and placed Uta, Urosaurus, Sator, and Sceloporus on one side and Uma, Holbrookia, and Callisau- rus on the other. Primitive to both groups was Petrosaurus. A sand lizard resur- rected by Clarke (1965) was Troschel's (1852) genus Cophosaurus (C. texanus, previously Holbrookia texana) .

Presch (1969) rejected Etheridge's re- moval of PJirynosoma from the scelopor- ines and placed Phrynosoma with the sand lizards as a primitive member of that group. On the basis of scleral ossicles, Presch (1970) indicated that Petrosaurus is a primitive member of the Sceloporus branch. Ballinger and Tinkle (1972) pro- posed an early separation of the Uta and Petrosaurus stock from the ancestor of Urosaurus, Sator, and Sceloporus.

Several characters suggest further modi- fication of the above arrangement. Our jiroposed phylogeny of sceloporine genera is illustrated in Figure 1. Urosaurus shows a tendency for enlarged scales near the midline of the dorsum. This trend is further developed in Sator, which has en- larged dorsals and granular laterals. The migration of enlarged scales around the sides of the body and the increase in scale size and degree of imbrication, mucrona- tion, and carination is a general trend along the chain of genera from Petrosau- rus to Sceloporus. The new phylogeny is also supported by the gradual decrease in development of the gular fold, which is completely lost in all species of Sceloporus in Group III. Most of the species in Group I have what Smith (1939) called a rudimentary gular fold. Some of the species in Group II show a less pro- nounced tendency to develop a gular fold, and Group III lacks it completely. The gradual loss of the gular fold in the Sceloporus complex is more probable than a loss (from Petrosaurus to Sceloporus) and subsequent redevelopment (from Sceloporus to Uta, Urosaurus, or Sator). This reversal of the phylogeny resolves a question raised by Smith (1946:178): "It is a curious fact that all genera that have sprung from Sceloporus have developed a gular fold including Sator, a Baja Cali- fornia genus. The tendency to develop this fold a})pears to be restricted to the ])rimitive groups of Sceloporus . . . and these are the groups from which Uta,

March 1975

LARSEN, TANNER: SCELOPORINE LIZARDS

Urosaurus, and Sator independently ap- pear to have been derived."

Although Smith pointed to this prob- lem, he nevertheless accepted Mittleman's arrangement of the sceloporine genera. More recently, Smith (per comm.): has agreed that Sceloporus ma}' be derived with respect to Uta^ Urosaurus. and Sator. This position has also been suggested by Hall (pers. comm.): "Inspection of the structure of the femoral pores and their surrounding scales, and the development of mucronation and carination of the body scales, to mention but two sets of charac- ters in various primitive Sceloporus and in other sceloporine genera, will suggest that Sceloporus is derived even in respect to Uta and Urosaurus. ''

We suggest the following conclusions with regard to the new phylogeny and published data on hip ratios of displaying males (Purdue and Carpenter, 1972a, 1972b). The hip ratio (vertical hip move- ment to vertical eye movement) increased from Petrosaurus (0.68) to Uta (average 0.74) to Urosaurus (average 1.06). After the transition from Sator (no published data on hip ratios) to Sceloporus, the trend reversed and hip ratios decreased from an average of 1.21 in Group I to 0.66 in Group II to 0.34 in Group III (averages computed from Purdue and Carpenter, 1972b).

Etheridge (1964) illustrated clavicles and scapulocoracoids of 8 sceloporine genera (excluding Phrynosoma) . If his drawings are superimposed on the new phylogeny (Fig. 2), two trends are ap- parent: (1) a gradual development of the scapular fenestra (top groove) from Petrosaurus to Sceloporus Group III, and (2) an increase in size of the clavicular hook. If Urosaurus and Uta were derived from Sceloporus, the scapular fenestra would have developed and then disap- peared from Petrosaurus to Sceloporus to Uta. This improbable reversal is similar to the problem with the gular fold. We are persuaded that the new phylogeny is more probable.

Intrageneric Phylogeny

The first ph^dogenetic schemes for the genus Sceloporus were proposed by Smith (1934, 1937a, 1937b, 1938, 1939). Other workers have recently modified the phy- logeny on the basis of karyology (Cole,

1970, 1971a, 1971b; Hall, 1971, 1973), and behavior (Bussjaeger, 1971).

Larsen and Tanner (1974) redefined relationships among the species in the genus Sceloporus. We used Ward's clus- ter analysis (Wishart, 1968) to cluster 55 species on the basis of external characters, cranial osteology, karyology, behavior, and zoogeography (Fig. 3). We then used step- wise discriminate analysis (Dixon 1967) and found that the arrangement of groups and subgroups is significant at the .999 level of confidence (Table 1).

Although Ward's cluster analysis pro- vides a phenetic dendogram, it does not give any indication as to which branch of a cluster is derived and which is primi- tive. In 1939 Smith said, "The most primitive form of this group is undoubted- ly lunaei which is closely related to for- mosus malachiticus'' (p. 60). In other words, lunaei is the most primitive form

PETROSAURUS

Fig. 2. Clavicles and scapulocoracoids of sev- eral sceloporines. All illustrations except Scelo- porus I, Sceloporus II, and Sceloporus III are from Etheridge (1964).

4

GREAT BASIN NATURALIST

Vol. 35, No. 1

Tabt.k 1. Groups

and subgroups in the genus Sceloporus

Group I (7 spp

)

Group II (20 spp.)

Group III (33 spp.)

Subgroup A ( 1

spp.)

Subgroup A (7 spp.)

Subgroup A (9 spp.)

gadoviae

grammicus

spinosus

Subgroup B (2

spp.)

pictus

orcutti

couchi

megalepidurus

clarki

memami

cryptus

melanorhinus

Subgroup C (4

spp.)

shannonorum *

magister

maculosus

heterolepis

olivaceus

parvus

asper

cautus

jalapae

Subgroup B (2 spp.)

horridus

ochotei-enae

pyrocephalus

edwardtaylori

nelsoni

Subgroup B (7 spp.)

Subgroup C (3 spp.)

formosus

scalaris

lunaei

goldmani*

nialachiticus

aeneus

acanthinus

Subgroup D (4 spp.)

Subgroup C (5 spp.)

siniferus

undulatus

carinatus

virgatus

utifonnis

woodi

squamosus

occidentalis

Subgroup E (4 spp.)

graciosus

variabilis

Subgroup D (4 spp.)

cozumelae

jarrovi

teapensis

lineolateralis

chrysostictus

ornatus dugesi Subgroup E (8 spp.) torquatus cyanogenys bulleri insignis* macdougalli mucronatus serrifer poinsetti

* Species not examined in this study.

ill the spinosus s])ecies group because it is most similar to a member of the next closest group (formosus). This statement by Smith is consistent with the following method of converting a phenetic dendro- gram into a phylogeny (Fig. 4): If "A" is primitive to "B" it is less derived from (more similar to) the stem species "G." The more primitive member of the other cluster ("C" or "D") will also be more similar to "G." The more primitive mem- bers of the two clusters will therefore be phylogenetically "closer" and phenotypi- cally more similar than any other com- bination from the two clusters. This rule can be applied objectively with a similar- ity matrix.

When all possible pairs between adja- cent clusters are compared, the two most similar species are considered jjrimitive within their res|)ective clusters. This technique will convert a dendrogram into a phylogeny.

Ward's cluster analysis and the above phylogeny techni(|ii(> were repeated sever-

al times using external and osteological characters, distribution, karyology, be- havior, and combinations of the above. (See Larsen and Tanner, 1974, for a pre- sentation of results.) The differences among results were resolved subjectively to produce a composite phylogeny (Fig. 5). This ])rocedure is based on several assumptions which are admittedly vul- nerable. To restrict the scope of our study it was assumed that the alpha taxonomy is complete and correct. That is, it was assvimed that all species of Sceloporus are now named and correctly defined in the literature. Of course, this assumption may be incorrect. But the purpose of our study is to produce a general overview and not a detailed taxonomic review. The de- tails near the ends of branches are there- fore tonlativo and stibject to future re- view.

In spite of the large number of charac- ters considered (over 80), these results are also subject to errors due to parallel- ism, convergence, varying rates of diver-

March 1975

LARSEN, TANNER: SCELOPORINE LIZARDS

Formosus __^ spinosus .^_^ Horndus _—

Olivaceus

Cqutus

Adieri

Molachiticus Luna«i

Lundalli ^^— Acantninus ~__ Edwprdtoylori.

Orcutti —^—

Magistcr

Undulatus

Occidentalis ^ Virgatus '— ^— Graciosus ^^— i Torquatus sernfer

Mucronatus

Cyonogenyi

Bulleri Poinsetti jarrovi -^—— Linaolateralis- Ornotus Dugesi ^—

Atper

Heterolepis_ Grammicus

Megolopidurus. Pictus ^^-^^^

Ochoterenae

Jalopae

scolons

Aeneus

pyrocephalus- Nelsoni - Melonorhinus. siniferus Connotui

Utiformis

Variabilis

Cozumela*

Teapensif '^^— Chrysostictus- squamosus^^ Parvus ^^.— Maculosus^— Couchi MerriamI— Cadoviae

^^

^

^

^

i

;=^

0.5

16

Fig. 3. Dendrogram generated by Ward's cluster analysis of external, diaracters (82 characters).

skull, and distribution

gence, pleiotrophy, and other cases in which the phenotype is not a direct mani- festation of the genotype. All phylogene- tic conclusions are subject to these liinita- tions, and the systematist can do little more than acknowledge the circumstantial nature of his evidence.

We propose that SceJoporus is derived from Uta through Urosaurus and Sator (see above). Smith (1938) suggested that tlie connection between these genera is from Urosaurus ornatus to Sceloporus couchi. Smith included couchi in the variabilis species group.

Figure 6 shows the arrangement of species in Smith's variabilis, maculosus, and mcrrianii groups according to Smith (1939, Fig. 42) and the new phylogeny. Four of these species {couchi, parvus, maculosus, and merriami) are transferred to Group I. Smith may have allowed for this by placing these four species on one side of his tree next to Uta. If Uta {Uta,

Urosaurus, and Sator) is considered primi- ti^'e to Sceloporus, then Smith's evidence supports our conclusion that Group I is primitive to the other two groups in Sceloporus. The remaining species in Smith's variabilis group {variabilis, coz- umelae, and teapensis) are placed in Group II.

Smith (1939:239) allowed for the re- moval of parvus and couchi from the var- iabilis grouj) with this statement:

That parvus and couchi are only dis- tantly related to the remainder of the group is shown by the widely different charac- ter of the ventral coloration in the males, smooth head scales, larger number of fem- oral pores, and general habitus. ... It is my belief that this section approaches more closely the ancestral stock of Uta than the other species of the variabilis group.

Smith (p. 239) also associated merriami with Uta: "It w^ould appear that merri- ami is closely related to Uta. and that Uta

GREAT BASIN NATURALIST

Vol. 35, No. 1

B

E

1

1

F

1

1 G

IF A-C = l

A-D=2

B-C = 2

B-D

= 3

THEN--

Fig. 4. Phylogeny theory- K the phenetic distance between "A" and "C" is less than that between any other pair, then "A" and "C" are primitive members in clusters "E" and "F."

arose from the forms now extinct which closed the present gap between couchi and merriami.'" Note that our new ar- rangement places merriami and couchi together.

Another divergence from Smith's phylo- genetic tree is the addition of chrysostic- tus to the variabilis group. Smith (p. 239) supports this inclusion (and the close proximity of the siniferus group) : "An- other group close!)' related to the variab- ilis section is the siniferus series, which closely approaches the variabilis group through cuprous. . . . The chrysostictus group is also closely related."

Thus it can be seen that Smith allowed for the possibility of removing parvus and couchi and adding chrysostictus, which changes his variabilis group into the new variabilis group.

Smith stated that the siniferus group "closely approaches the variabilis group" and yet his illustration (1939, Fig. 3) has these groups separated by several other groups. In the new phylogeny they are adjacent.

Figure 7 compares Smith's arrange- ment of his chrysostictus. utiformis and siniferus groups with the new arrange- ment of the same species. Besides the placing of chrysostictus in the variabilis group (which has already been ex- plained), the only major difference in Figure 4 is the removal of ochoterenae to place it in Group I. (The inclusion of utiformis in the siniferus group is mi- nor) . Smith listed 1 1 diagnostic characters of the siniferus group. In three cases he said "except ochoterenae'' and in another "except ochoterenae and cupreusT He (p. 301 ) said, "Postanals tending to be poorly developed (except ochoterenae and cupreus); two postrostrals (except ocho- terenae, without postrostrals) ; . . . ventral scales pointed or, at least not notched (ex- cept ochoterenae in which they are notched) . . . males without distinctive ventral coloration (except ochoterenae).'"

If size is discounted, then ochoterenae is different in 4 of the 10 diagnostic char- acters for the siniferus group. S. ocho- terenae also has more femoral pores than any other species in Smith's siniferus group. Smith's conclusions, therefore, would not be seriously challenged if ochoterenae were removed from the sinif- erus group and placed in Group I next to jalapae. In fact, when describing ocho- terenae., Smith (p. 309) said, "three or four scales on anterior border of ear, not so large as in jalapae.'' So apparently he was comparing these two species.

Smith included jalapae in his scalaris group, which is otherwise identical to the new scalaris group (Fig. 5). Removing jalapae from the scalaris group to place it in the primitive Group I is supported by the following statement b^- Smith (p. 331):

The only species doubtfully inchuled in this group is jalapae, which differs from the remaining fonns in having lateral scales in distinctly oblique rows, and in lacking postrostrals [as does ochoterenae]. . . .

5. jalapae is clearly, the most primitive member of the group. S. scalaris, aeneus and goldmani are clearly more closely re- lated to each other than any one of these is to jalapae.

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LARSEN, TANNER: SCELOPORINE LIZARDS

HORRIDUS

MELANORHINUS

MALACHITICUS

ACANTHINUS

OCCIDENTALIS

SCALARIS I UTirORMIS

GOLDMANI

VARIABILIS

TEAPENSIS

Fig. 5. Proposed phylogeny for the genus Sceloporus. (* = species not examined.)

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Vol. 35, No. 1

PARVUS MACULOSUS MERRIAMI

TEAPENSIS COZUMELAE

Fig. 6. Phylogeny of Smith's (1939) variabilis, maculosus. and merriami groups according to Smith (A) and the new phylogeny (B).

CHRYSOSTICTUS

VARIABILIS GROUP

SCALARIS GROUP

OGHOTERENAE

CHRYSOSTIGTUS

VARIABILIS GROUP

SCALARIS GROUP

OGHOTERENAE

Fig. 7. Phylogeny of Smith's (1939) chrysostictus, utiformis, and siniferus groups according to Smith (A) and the new phylogeny (B).

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LARSEN. TANNER: SCELOPORINE LIZARDS

(iroup I includes: parvus, couchi, ma- ( ulosus, mcrriami, ochoterenat\ jalapae, aiul gadoviac. the most primitive. Smith ( p. 362) inchided gadoviac with nelsoni and pyrocephalus in the pyrocephalus group. But once again he outhned rea- sons why gadoviac could be removed and [)laced in Group I. "5. gadoviae differs widely from other members of the group in having very small dorsal scales, a large number of femoral pores, a postfemoral dermal pocket, very small scales on pos- terior surface of the thighs, and many other minor characters." S. gadoviae is also the only member of this group to have a vestigial gular fold as mentioned by Smith (p. 374): "scales immediately preceding gidar fold region somewhat re- duced in size." All of these characters are diagnostic of Group I, and this primi- tive placement is therefore natural. In fact. Smith (p. 363) said, "I assume gadoviae to be nearest the primitive type, as it retains certain characters of the variabilis group, from which I believe it was derived."

The main character on which Smith (p. 363) based his inclusion of gadoviae with the pyrocephalus group is the strong com- pression of the tail: "That the group is a natural one is more or less assured by its compact range and by the common char- acter of the compressed tail, which is otherwise unknown in the genus." In view of the many characters supporting the placement of gadoviac in Group I, we propose that a compressed tail developed twice: once in the pyrocephalus group, and once in gadoviae. Smith (p. 363) gave further support to this placement of gadoviae: "The assumption that gadoviac is a remnant of a primitive stock is sup- ported by its secretive habits and its re- striction to a somewhat arid region."

The most serious difference between the new phylogen^- and that of Smith is the placement of the gramniicus and me- galepidurus groups. In both phylogenies the species are arranged in a similar man- ner within these groups. But Smith placed these groups next to the jormosus group with the large-scaled, large-sized species, and we ha\e moved them to a primitive position in Group II. How- ever, we propose that the grammicus group (we have combined Smith's gram- micus and hetcrolcpis groups) is the most primitive in Group II. In fact, Smith

(1938:552) said "the microlepidurus [our grammicus^ group is assumed to be the most primitive of these [the large-scaled, large-sized sjiecies], largely because of its very small scales." This greater separ- ation between the grammicus and jormos- us groups is further justified by the fact that the diploid number of chromosomes is 22 (derived) in the jormosus group and 32 (primitive) in the grammicus group. We propose, therefore, that some of, the similarities between grammicus and jor- mosus (coloration, dorsal-scale count, ovo- viviparity, and preference for an arboreal habitat) are a result of convergence as is true of gadoviae and the pyrocephalus group.

The only remaining difference from Smith's jormosus group is his inclusion of asper, which we have moved to the gram- micus group. This move is justified by the fact that asper has 32 chromosomes, as do the other members of the grammicus group. If the grammicus grou]:) is re- moved from Smith's large-scaled, large- sized branch, the remaining species are the same as those included in Group III. This grouping (the omission of grammi- cus) was allowed by Smith (1938:552):

The relatively small size of the species of the undulatus group must be assumed as a parallel development rather than a direct inlieritence of the small size of the ancestor in the variabilis group, for the close rela- tionship of the spinosus and undulatus groups cannot logically be disputed, nor is the close relationship of the spinosus, lor- qualus and formosus groups doubtful."

Smith and Taylor (1950) included the following species within the undulatus group: undulatus, cautus, occidentalism and woodi. Since then, virgatus has been raised from subspecific to specific status (Cole, 1963). Smith (1939) placed fjrac/- osus adjacent to the undulatus group, so the only discrepanc}' between the two classifications is the placement of cautus, which we have moved to the spinosus group next to olivaceus. This mo^'ement is justified by the fact that there is a zone of intergradation between cautus and oli- vaceus (Hall, pers. comm.).

Bussjaeger (1971:151) remarked:

The relation of cautus and olivaceus and the undulatus group of Sceloporus has been questioned. Hall's data indicated that these two species were the same and limited data on their displays indicate that they are similar. If one accepts that they are syn-

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onyms, then olivaceus (cautus) would be the connecting link between the spinosus and undulatus groups.

However, rather than use these forms as a link between species groups, we have placed them together in the spijiosus group.

Smith (1938:554) indicated that the torquatus group consited of 2 subgroups: "It appears that soon after the separation of the torquatus stock from the other groups of Sceloporus, there was a separa- tion into two divisions, one of which ex- hibited a tendency to develop small scales, the other large scales." We have recog- nized the small-scaled division as the jarrovii group.

Figure 8 shows the phylogeny of the jarrovii group according to Smith (1938, Fig. 4) and the new arrangement. Al- though he placed lineolateralis further away from jarrovii in his diagram. Smith (p. 556) did say, "S". jarrovii appears to be most closely related to lineolateralis. From this species, or its ancestors, the re- maining species of the small-scaled divi- sion have obviously been derived."

Figure 9 shows the phylogeny of the torquatus group according to Smith (1938,

LINEOLATERALIS

Fig. 8. Phylogeny of jarrovi group according to Smith (1938) (A) and the new phylogeny (B).

Figs. 3-4) and the new arrangement. There seems to be little similarity here, except that torquatus is derived from serrifer, and poinsetti is derived from cyanogenys in both trees. Smith (1938: 555) raised a question about the ancestral position of serrifer:

S. serrifer appears to be the oldest of the large-scaled species. The postulation that this species, which is one of the larger ones

POINSETTI

CYANOGENYS

Fig. 9 Phylogeny of torquatus group according to Smith (1938) (A) and the new phylogeny (B).

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LARSEN, TANNER: SCELOPORINE LIZARDS

11

of the genus, and one having large scales, is nearest to the ancestral type of the large- scaled division of the torquatus group may appear to be contradictory to the postula- tion that Sceloporus is derived from small species with small scales. However, my as- sumption seems to be justified by the fact that serrifer occupies a southern position on the periphery of the geographical area now occupied by the torquatus group.

The reason for this paradox is that Smith assumed speciation in Group III was from south to north. The data in 1938 strongly supported this conclusion. Obviously, Smith did not believe that a peripheral location is necessarily primitive, because on the next page (556) he said, "S". mu- cronatus appears to be the nearest to the ancestral type of these three species {cy- anogenys, poinsetti and omiltemanus) de- spite the fact that it has larger scales than they. I so conclude because of its central- ized geographical position with relation to the area occupied by the other three forms."

So the basic problems can be solved, and the trend is indeed from small to large size and small to large scales if this group was developed from north to south rather than south to north. Smith indi- cated a northward development from ser- rifer to torquatus to mucronatus to cyano-

genys, and our phylogeny indicates a southward development from cyanogenys to mucronatus to serrifer to torquatus. An ancestral placement of cyanogenys is fur- ther supported by Smith (1939:209): "Species of this group are as a rule con- fined to rocky habitats. So far as I am aware, only cyanogenys tends to live on or near the ground." Thus, the new ]:)hylogeny indicates a trend in this group from small-sized, small-scaled ground dwellers to large-sized, large-scaled rock dwellers. With this reversal in direction, the remaining differences between the two phylogenies in Figtire 9 are negligible and the trends within this group fit the overall phylogeny of the genus.

In the genus Sceloporus, the spinosus group has been the object of more system- atic study than any other. No less than four different phylogenetic trees have been proposed by Smith, Bussjaeger, Cole, and Hall. The confusion is further compound- ed by the fact that the spinosus group is the largest in number of species and sub- species. The four phylogenetic trees and our conclusions are presented in Figure 10. Smith (1939) included acanthinus, lunaei. and lundelli with this group. In 1950, he and Tavlor moved acanthinus

Fig. 10. Phylogeny of spinosus group according to Smith (1939), Cole (1970), Bussjaeger (1971), Hall (pers. comm. 1973), and the new phylogeny (L and T).

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and lunaei into the formosus group. How- ever, in 1939 Smith (p. 60) said, "The most primitive form of the group is un- doubtedly lunaei. which is closely related to formosus malachiticus. S. acanthinus is a near relative of lunaei. as is also lun- delli.'" It should therefore be acceptable to remove lundelli from the spinosus group and place it in the formosus group next to lunaei as we have done.

Behavioral data also support this ar- rangement. Bussjaeger (1971:136) ob- served:

The display-action-patterns of lundelli gaigei of the spinosus group and asper, acan- thinus acanthinus and a. lunaei of the formosus group were quite similar with peaked single units and multiple units. Sceloporus asper and lundelli seemed to share more elements.

In his conclusions, Bussjaeger (p. 151) an- ticipated the new position of S. lundelli:

The status of lundelli is questionable. . . . Its display-action-pattern was between acan- thinus and orcuiti; but the pattern was based on only one female. More data are needed to establish this species relationship. At present it should be left in the spinosus group, although it appears to be closer to the formosus group.

Cole's (1970) phylogenetic tree would xiot allow the removal of lundelli from this group unless melanorhinus and clarki were placed elsewhere. Cole (p. 39, Fig. 17) showed how four centric fusions could change the melanorhinus-clarki karyotype into the typical pattern for this group. According to Cole's assumption that only fusions (i.e., no fissions) are possible, melanorhinus and clarki are primitive not only for this group, but also for the genus Sceloporus. and for the entire fam- ily Iguanidae! As demonstrated by Web- ster, Hall, and Williams (1972), chromo- somal evolution can occur by fission as well as fusion. We believe this is the only acceptable explanation for the karyo- type in melanorhinus and clarki. If fission is accepted as well as fusion, Cole's data provide support for our arrangement of orcutti, clarki. and melanorhinus. (They also confirm the primitive position of lundelli and permit its placement in the formosus group.)

If clarki and melanorhinus are derived