TABLE 1

CYTOSYSTEMATICS OF SCELOPORUS

Smith (1939) species groups and species according to Smith and Taylor (1950)1 2n karyotype formula2 sources3

SMALL-SIZED, SMALL-SCALED SPECIES
VARIABILIS
couchi 2n=34 (10MM,2SM,20m,xx♀ xy♂) H2
parvus 2n=34 (10MM,2SM,20m,xx♀ xy♂) H2
variabilis 2n=34 (10MM,2SM,20m,xx♀ xy♂) H2
teapensis 2n=34 (10MM,2SM,20m,xx♀ xy♂) H2
cozumelae ? - -
MACULOSUS
maculosus4 2n♀=34
2n♂=33		 (10MM,2SM,18m,xx,x1x1)
(10MM,2SM,18m,xx,x1y1)		 C4
maculosus5 2nª40 A
MERRIAMI
merriami 2n=46 (24AM,20m,xx♀ xy♂) C4,H2
CHRYSOSTICTUS
chrysostictus 2n=34 (10MM,2SM,20m,xx♀ xy♂) C4
SINIFERUS
siniferus 2n=34 (10MM,2SM,20m,xx♀ xy♂) H2
squamosus ? - -
carinatus ? - -
ochoterenai ? - -
UTIFORMIS
utiformis 2n=34 (10MM,2SM,20m,xx♀ xy♂) C4,H2
SCALARIS
jalapae 2n=34 (10MM,2SM,20m,xx♀ xy♂) H2
goldmani ? - -
scalaris 2n=24 (10MM,2SM,10m, x2x2♀ x2y2♂) L1,H2
aeneus 2n=24 (10MM,2SM,10m, x2x2♀ x2y2♂) H2
PYROCEPHALUS
gadovae 2n=34 (10MM,2SM,20m,xx♀ xy♂) C5,H2
nelsoni 2n=34 (10MM,2SM,20m,xx♀ xy♂) C5,H2
pyrocephalus 2n=34 (8MM,2SM,2SAM,20m, xx♀ xy♂) C5,H2

LARGE-SIZED, LARGE-SCALED SPECIES
SPINOSUS
orcutti orcutti 2n=34 (10MM,2SM,20m,xx♀ xy♂) C3,H2
[(orcutti) licki 2n=34 (10MM,2SM,20m,xx♀ xy♂) H2
[n. sp.] 2n=34 (10MM,2SM,20m,xx♀ xy♂) H2
clarki 2n=40 (2SM,2MM,16AM,18m, x3x3♀ x3y3♂)
[note polymorphisms for enlargement of micro-autosome, inversion of an AM]		 L2,C3,H2
melanorhinus 2n♀=40
2n♂=39		 (2SM,2MM,14AM,XX,18m, x3x3)
(2SM,2MM,14AM,XY,18m,x3)
[note polymorphisms for enlargement of micro-autosome, probably same as clarki polymorphism		 C3,H2
[(magister) zosteromus] 2n=30 (10MM,2SMM,18m - xy indistinguishable) H2
magister (mainland) 2m=26 (8MM,2SMM,2SAM,14m - xy indistinguishable) L2,C3,H2
horridus 2n=22 (10MM,2SMM,10m - xy indistinguishable) C3,H2
spinosus (10MM,2SMM,10m - xy indistinguishable) C3,H2
edwardtaylori (10MM,2SMM,10m - xy indistinguishable) C3,H2
lundelli (10MM,2SMM,10m - xy indistinguishable) C3
olivaceus6 (10MM,2SMM,10m - xy indistinguishable) C3,H2
UNDULATUS
cautus6 2n=22 (10MM,2SMM,10m - xy indistinguishable) C6,H2
exsul7 ? - -
undulatus 2n=22 (10MM,2SMM,10m - xy indistinguishable) C6,H2
occidentalis 2n=22 (10MM,2SMM,10m - xy indistinguishable) C1,C6,J,H2
virgatus8 2n=22 (10MM,2SMM,10m - xy indistinguishable) C2,C6
woodi 2n=22 (10MM,2SMM,10m - xy indistinguishable) C2,H2
FORMOSUS
formosus 2n=22 (10MM,2SMM,10m - xy indistinguishable) H2
adleri9 2n=22 (10MM,2SMM,10m - xy indistinguishable) H3
tanneri10 ? - -
stejnegeri ? - -
lunae ? - -
salvini11 ? - -
smaragdinus11 ? - -
taeniocnemis11 ? - -
internasalis11 ? - -
acanthinus11 ? - -
malachiticus 2n=22 (10MM,2SMM,10m - xy indistinguishable) H2
asper12 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note fixation of enlargement of microautosome, probably same is involved in clarki-melanorhinus polymorphism]		 H2
GRACIOSUS
graciosus (10MM,2SMM,18m - xy indistinguishable)
[note same as zosteromus except sattelite]		 C4,C7,H2
GRAMMICUS
heterolepis 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 H2
shannonorum13 2n♀=32
2n♂=31		 [same as heterolepis] H2
grammicus [S] 2n♀=32
2n♂=31		 [same as heterolepis] H1,H2
grammicus [P1] 2n♀=32-34
2n♂=31-33		 [same as heterolepis but polymorphic for fission MM pair 1] H1,H2
grammicus [F6] 2n♀=34
2n♂=33		 [same as heterolepis except fixed for fission MM pair 6] H1,H2
grammicus [F5] 2n♀=34
2n♂=33		 [same as heterolepis except fixed for fission MM pair 5] H2
grammicus [F5+6] 2n♀=36
2n♂=35		 [same as heterolepis except fixed for fissions MM pairs 5 and 6] H2
grammicus [FM1] 2n♀=40-44
2n♂=39-43		 [same as heterolepis except fixed for fissions MM pairs 2, 4, 5 and 6; polymorphic for fissions of MM pairs 1 and 3] H2
grammicus [FM2] 2n♀=44-46
2n♂=43-45		 [same as heterolepis except fixed for fissions MM pairs 1, 2, 4, 5 and 6 plus microchromosome pair 14; polymorphic for fission of MM pair 3] H2
MEGALEPIDURUS
megalepidurus14 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note fixation of enlargement of microautosome, probably same is involved in clarki-melanorhinus polymorphism]		 H2
subpictus15 ? - -
cryptus16 2n=22 (10MM,2SMM,10m - xy indistinguishable) H2
TORQUATUS
dugesi 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 H2
ornatus 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 H2
jarrovi17 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 C1,H2
mucronatus 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 H2
bulleri 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 H2
insignis18 ? - -
torquatus 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 H2
poinsetti 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 C1,H2
cyanogenys 2n♀=32
2n♂=31		 (10MM,2SM,16m,x4x4, x3x3)
(10MM,2SM,16m,x4x3y4)
[note same as asper without enlargement of microautosome]		 H2
serrifer19 ? - -
macdougalli20 ? - -
  1. Where presently valid taxonomic usage differs from the Smith and Taylor (1950) standard, the species name will be referenced to a footnote below which cites the author(s) of the revision. Complete references will be found in the bibliography. See also Smith and Smith (1976). Species which will result from my intended taxonomic revisions are enclosed in brackets.
  2. MM = metacentric macrochromosome,- SMM = submetacentric macrochromosome, SAM = subacrocentric liiacrochromosome, AM = acrocentric macrochromosome, m = micro-autosome. Sex chromosomes indicated by small letters are microchromosomes, while those indicated by caps are macrochromosomes. Subscripts on sex chromosomes indicate probably different derivations. The simple xy designates the primitive sceloporine sex chromosomes (Pennock, et al., 1969). Karyotype forumulas do not indicate several minor inversion differences, translocations, or polymorphisms noted in Cole's papers.
  3. Sources are coded as follows (complete references will be found in the bibliography): A = Carol Axtell, pers. comm. - her slides were checked by me; C1 = Cole, Lowe, and Wright, 1967; C2 Cole and Lowe, 1968; C3 = Cole, 1970; C4 = Cole, 197la; C5 = Cole, 1971b; C6 = Cole, 1972; C7 = Cole, 1975; H. = Hall . and Selander, 1973; H = Hall, 1973 - photographs and localities are presented for clarki, melanorhinus, asper, megalepidurus, the grammicus group and the torquatus group. Only chromosome formulas are listed for the other species. It is planned to publish all of these data in the open literature; H3 = Hall, unpub.; J = Jackson and Hunsaker, 1970; L1 = Lowe, Wright, and Cole, 1966; L2 = Lowe, Cole, and Patton, 1967.
  4. Specimens from southern Coahuila (Cole, 1971a).
  5. Specimens from near Pedrecena, Durango - the type locality for maculosus (C. Axtell, pers. comm.).
  6. My collections include probable intergrades between olivaceus and cautus involving the character Smith (1939) uses to distinguish the undulatus and spinosus species groups.
  7. Dixon, et al., 1972.
  8. Cole, 1963.
  9. Smith and Savitzky, 1974.
  10. Smith and Larsen, 1975.
  11. Stuart, 1971.
  12. S. asper does not belong in the formosus group. Based on karyotypes it clearly is placed with megalepidurus and the grammicus and torquatus groups (Hall, 1973).
  13. Langbartel, 1959 - I do not accept Webb's (1969) lumping of shannonorum with heterolepis 
    1. because the two species are absolutely separated by the climatic barriers of the deep barrancas of the Rio Grande de Santiago drainage of Lake Chapala and 
    2. because the conspicuously enlarged paravertebral scales that so obviously distinguish heterolepis are not found in any shannonorum population. The fact that some heterolepis "intergrades" have intermediate dorsal midline scale counts is completely irrelevant. These two species are certainly more distinct from one another than many recently discovered Sceloporus that prove their genetic isolation in sympatry with one another.
  14. Dassmann and Smith (1974) lump pictus with megalepidurus, based on my discovery of intergrading populations.
  15. Lynch and Smith, 1965 - this species should probably be reallocated to the 2n = 22 formosus group.
  16. Smith and Lynch, 1967 - cryptus obviously belongs with the 2n = 22 formosus group.
  17. Webb and Hensley (1959) lump lineolateralis with jarrovi.
  18. Webb, 1967.
  19. Stuart (1970) lumps prezygus with serrifer.
  20. Smith and Bumzahem, 1953.