NOTES
Caribbean Journal of Science, Vol. 29, No. 3-4, 250-253, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
The Genetic Relations of
Anolis cristatellus
(Sauria: Polychridae) from
Hispaniola and Puerto Rico
PETER A. ZANI 1,3 AND SHELDON I. GUTTMAN ,1 1Department
of Zoology, Miami University, Oxford, Ohio 45056.
ROBERT POWELL ,2 2Department of Natural Sciences, Avila
College, Kansas City, Missouri 64145.
Anolis cristatellus, a common lizard of the Puerto
Rican Bank (German et al., 1980b), has been introduced into the vicinity of La Romana, Republica Dominican, and were well established when first noted
by Grant (1956). Williams (1977) suggested that these
animals may have been introduced when a sugar mill
was built by a Puerto Rican based company during
the years 1910-1920. These lizards are firmly entrenched in and around the city and have nearly displaced endemic A. cybotes from these areas (Fitch et
al., 1989).
German et al. (1980b) attempted to reconstruct the
phylogeny and colonization sequence of A. cristatellus, but failed to include representatives from Hispaniola. The objective of our study was to compare
the genetic characteristics of two representative populations from Puerto Rico with that on Hispaniola.
Anolis cybotes from Hispaniola was examined to elucidate interspecific differentiation. This species was
once considered a close relative of cristatellus (Williams, 1976), and although no longer the case (Gor-
3
Present address: Department of Zoology, University of Oklahoma, Norman, Oklahoma 73019.
man et al., 1980a; German et al., 1983; Burnell and
Hedges, 1990), they are ecological equivalents on Hispaniola (Fitch et al., 1989).
Anoles were collected in the Dominican Republic
(DR) during June 1991 (30 A. cristatellus from La Romana and 29 A. cybotes from near Santo Domingo),
transported alive to Kansas City, frozen, and shipped
to Miami University on dry ice. Puerto Rican (PR)
specimens were Obtained from Rio Piedras (n = 13)
and Mayaguez (n = 27) and shipped alive to Miami
University. All were stored at –70°C until tissues were
processed. Specimens were accessioned into the vertebrate collection at Miami University (lots R155R159).
Heart, liver, kidney, and muscle tissue samples were
dissected from the frozen animals, homogenized in
two volumes of buffer (Yang et al., 1974) and centrifuged at 14,000 rpm for 40 min. The supernatant was
removed and frozen at –70°C. Horizontal starch gel
electrophoresis was conducted following procedures
outlined by Selander et al. (1971), McKinney et al.
(1972), and Buth et al, (1980), with the following modifications, After comparing resolution of loci on Triscitrate pH 7.0 to that with the Clayton and Tretiak
(1972) buffer system, we found that the latter gave
superior results for certain loci. Analysis of data was
carried out using the BIOSYS-I computer package of
Swofford and Selander (1981).
Individual enzyme loci examined and electrophoretic conditions appear in Table 1. Allele frequencies
and Chi-square results are presented in Table 2. A
matrix of genetic distance (Nei, 1978) and identity
estimates, along with mean heterozygosities are in
Table 3. Figure 1 is a phenogram based on the distance
values.
A Z-test was applied to the values for mean heterozygosity; these were not significantly different between the four populations. That heterozygosity was
not significantly reduced in the DR population suggests the lack of a founder effect. A large initial population or multiple introductions over time may be
responsible.
TABLE 1. Buffer systems used and presumptive gene loci scored. A: Tris-citrate pH 7.0 (Whitt, 1970); B:
Tris-citrate pH 8.0 (Selander et al., 1971); C: Tris-hydrochloric acid (Selander et al., 1971); D: Clayton-Tretiak
(1972); E: “Poulik” system (Selander et al., 1971).
Protein
Loci
Enzyme
commission
number
Esterase
Glucose-6-phosphate isomerase
Isocitrate dehydrogenase
Lactate dehydrogenase
Malate dehydrogenase
Nonspecific protein
Phosphoglucomutase
Phosphogluconate dehydrogenase
EST-1,2
GPI-1,2
IDH-1,2
LDH-1,2
MDH-1,2
NSP-1,2,3
PGM
PGDH
—
5.3.1.9
1.1.1.42
1.1.1.27
1.1.1.37
—
2.7.5.1
1.1.1.44
250
Buffer
system
C
D
D
A
D
E
B
B
NOTES
TABLE 2. Allele frequencies of Anolis populations.
Population 1. A. cristatellus from La Romana, Dominican Republic. 2. A. cristatellus from Mayaguez, Puerto
Rico. 3. A. cristatellus from Rio Piedras, Puerto Rico.
4. A. cybotes from Santo Domingo, Dominican Republic. Letters over allele frequencies represent results
from pairwise Chi-square tests between the four populations. The same letter over two populations indicates frequencies were not significantly different. Different letters between two populations indicate a
significant difference.
Population
Locus
1
2
3
4
a
b
y
0.55
0.45
xy
0.33
0.67
xy
0.27
0.73
x
0.29
0.71
x
1.00
x
1.00
x
0.96
0.04
x
1.00
x
x
x
1.00
1.00
1.00
y
0.59
0.38
0.03
x
x
1.00
1.00
x
0.08
0.92
y
0.85
0.16
x
1.00
y
0.28
0.72
x
0.92
0.08
x
1.00
x
1.00
x
1.00
x
0.96
0.04
x
0.95
0.05
x
x
0.19
x
0.27
1.00
0.82
0.73
y
0.41
0.55
0.03
x
x
x
1.00
1.00
1.00
y
x
0.07
0.93
0.98
0.02
x
0.04
0.92
0.04
x
1,00
x
0.98
0.02
x
1.00
x
1.00
x
0.37
0.63
x
0.46
0.54
x
0.27
0.73
y
1.00
x
x
x
y
0.09
EST-2
a
b
GPI-1
a
b
c
GPI-2
a
b
IDH-1
a
b
IDH-2
a
b
LDH-1
a
b
c
LDH-2
a
b
MDH-1
a
b
c
MDH-2
a
b
PGM
a
b
PGDH
a
T ABLE 2. Continued
Population
Locus
y
1.00
x
1,00
3
4
1.00
1.00
0.90
0.02
x
1.00
x
1.00
x
1.00
x
1.00
x
1.00
x
1.00
x
1.00
x
x
x
y
1.00
1.00
1.00
1.00
Allele
1
b
c
0.05
0.95
a
x
1.00
a
NSP-1
NSP-2
NSP-3
Allele
EST-1
251
a
b
2
Chi-square tests were also performed for each locus
examined (Table 2), For the two PR populations a
significant difference existed only at one of 15 loci,
These results confirm the close relationship of populations found on opposite ends of the island (although neither sample represented the southern
Puerto Rican morph, R. Thomas, pers. comm.). One
and two loci were found to be significantly different
between the La Romana/Rio Piedras and La Romana/
Mayaguez populations, respectively, Significant differences between A. cybotes and the DR A. cristatellus
existed in nine of the 15 loci examined. Comparisons
of A. cybotes with A. cristatellus from Mayaguez (8) and
Rio Piedras (7) yielded fewer significant differences.
Genetic distance between the Mayaguez and the
Hispaniolan samples was approximately twice (D =
0.111) that between the Rio Piedras and La Romana
(D = 0.069) populations. Genetic identity measurements offered similar results (Table 3). These data
suggest a closer affinity between anoles from La Romana and Rio Piedras than between the DR and Mayaguez populations.
Our results generally resemble those of German et
al. (1980b), except for the MDH-1 locus (Table 2). We
reexamined our animals for MDH on the same buffer
system used by German et al. (1980b) and our patterns
were unchanged. German et al, (1980b) found MDH
to be polymorphic in only one population on Anegada in the British Virgin Islands. We found MDH
to be polymorphic in all three populations of A. cristatellus examined. In addition, the MDH-1c was the
predominant allele in the Hispaniolan population,
while it was relatively scarce in the Puerto Rican populations. We also found a third allele (MDH-1a), undescribed by German et al. (1980b), in the population
from Rio Piedras.
We cannot explain why the c allele, present in very
low proportions in other populations (if at all), occurs
in such high frequency in the DR. Smith et al. (1983)
documented that different MDH alleles are selected
in Gambusia under different thermal conditions, Possibly an undetected stressor, such as temperature, is
acting on the Hispaniolan populations in a manner
selecting for MDH-1c. A biotic stressor may be the
competition with A. cybotes, which does not occur on
Puerto Rico.
NOTES
252
TABLE 3. Comparison of Nei’s (1978) genetic similarity coefficients (I) above diagonal and genetic distance
coefficients (D) below diagonal calculated between all pairs of populations. In addition, mean observed
heterozygosities plus standard error appear along the diagonal for each population. Populations as in Table 2.
Population
1
2
3
4
1
2
3
4
(0.094 ± 0.053)
0.111
0.069
0.580
0.895
(0.123 ± 0.059)
0.033
0.509
0.933
0.968
(0.129 ± 0.051)
0.451
0.560
0.601
0.637
(0.122 ± 0.047)
FIG. 1. A phenogram based upon the genetic distance values in Table 3. Populations are the same as in
Table 2.
Acknowledgments. — We wish to thank H. Bui, T.
Fobes, A. Lathrop, J. Lynxwiler, J. Moster, J. Parmerlee, Jr., P. Schell, M. Schreiber, D. Smith, J. Smith,
and L. White for their help in collecting DR specimens. Jose Ottenwalder of the Parque Zoologico National facilitated opportunities for field work. R. Thomas, University of Puerto Rico-Rio Piedras, and A.
Lewis, University of Puerto Rico-Mayaguez, graciously supplied animals from Puerto Rico. Permits were
provided by E. Bautista M., Director, Departmento de
Vida Silvestre, Republica Dominicana, and E. Cardona, Department of Natural Resources, Puerto Rico.
Preparation of animals was facilitated by M. Zehnder
and R. House and S. Fore aided in statistical analysis.
This investigation was supported in part by the Department of Zoology at Miami University and NSF
BBS-9100410 awarded to RP.
LITERATURE CITED
Burnell, K. L., and S. B. Hedges. 1990. Relationships
of West Indian Anolis (Sauria: Iguanidae): an approach using slow-evolving protein loci. Carib. J.
Sci. 26:7-30.
Buth, D. G., G. C. German, and C. S. Lieb. 1980.
Genetic divergence between Anolis carolinensis and
its Cuban progenitor, Anolis porcatus. J. Herpetol.
14:279-284.
Clayton, J. W., and D. N. Tretiak. 1972. Amine-citrate buffers for pH control in starch gel electrophoresis. J. Fish. Res. Bd. Can. 29:1169-1172.
Fitch, H. S., R. W. Henderson, and H. Guarisco. 1989.
Aspects of the ecology of an introduced anole:
Anolis cristatellus in the Dominican Republic. Amphibia-Reptilia 10:307-320.
German, G. C., D. G. Buth, and J. S. Wyles. 1980a.
Anolis lizards of the eastern Caribbean: a case study
in evolution. III. A cladistic analysis of albumin
immunological data, and the definition of species
groups. Syst. Zool. 29:143-158.
———, ———, M. Soule, and S. Y. Yang. 1980b. The
relationships of the Anolis cristatellus species group:
electrophoretic analysis. J. Herpetol. 14:269-278.
———, ———, ———, and ———. 1983. The relationships of the Puerto Rican Anolis: electrophoretic and karyotypic studies. In A. G. J. Rhodin and
K. Miyata (eds.), Advances in herpetology and
evolutionary biology, pp. 626–642. Museum of
Comparative Zoology, Harvard University, Cambridge, Massachusetts.
Grant, C. 1956. Report on a collection of Hispaniolan
reptiles. Herpetological 12:85-90.
McKinney, C. O., R. K. Selander, W. E. Johnson, and
S. Y. Yang. 1972. Genetic variation in the sideblotched lizard (Uta stansburiana). Studies in Genetics VII. Univ. Texas Publ. 7213:307-318.
Nei, M. 1978. Estimation of average heterozygosity
and genetic distance from a small number of individuals. Genetics 89:583-590.
Selander, R. K., M. H. Smith, S. Y. Yang, W. E. Johnson, and J. B. Gentry. 1971. Biochemical polymorphism and systematic in the genus Peromyscus. I. Variation in the old-field mouse (Peromyscus
polionotus). Studies in Genetics VI. Univ. of Texas
Publ. 7103:49-90.
Smith, M. H., M. W. Smith, S. C. Scott, E. H. Liu, and
NOTES
J. C. Jones. 1983, Rapid evolution in a post-thermal environment. Copeia 1983:193-197.
Swofford, D. L., and R. K. Selander. 1981. BIOSYS1: a FORTRAN program for the comprehensive
analysis of electrophoretic data in population genetics and systematic. J. Hered. 72:281-283.
Whitt, G. S. 1970. Developmental genetics of the
lactate dehydrogenase isozymes of fish. J. Exp. Zool.
175:1-35.
Williams, E. E. 1976. West Indian anoles: a taxonomic and evolutionary summary. I. Introduction and
species list. Breviora (440):1-21.
———. 1977. Anoles out of place: introduced anoles.
In E. E. Williams (ed.), The third Anolis Newsletter,
pp. 110-118. Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts.
Yang, S. Y., M. Soule, and G. C. German. 1974. Anolis
lizards of the eastern Caribbean: a case study in
evolution. I. Genetic relationships, phylogeny, and
colonization sequence of the roquet group. Syst.
Zool. 23:387-399.
Caribbean Journal of Science, Vol. 29, No. 3-4, 253-254, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
Notes on the Diets of
West Indian Liophis
(Serpentes: Colubridae)
R OBERT W. HENDERSON AND R OBERT W. BOURGEOIS ,
Section of Vertebrate Zoology, Milwaukee Public Museum,
800 W. Wells St., Milwaukee, Wisconsin 53233-1478.
The neotropical colubrid snake genus Liophis has a
wide mainland distribution (about 35 species from
Costa Rica to central Argentina; Dixon, 1989). Additionally, four species are endemic to the West Indies:
L. cursor (Martinique and Rocher du Diamant), L. juliae
(Guadaloupe, Marie-Galante, and Dominica), L. perfuscus (Barbados) and L. ornatus (St. Lucia and Maria
Major) (Schwartz and Henderson, 1991). Two dubious
records of a fifth species, L. melanotus, occurring on
Grenada are probably dismissible (Henderson, 1992),
but it has an extensive mainland distribution and also
occurs on Trinidad and Tobago (Dixon and Michaud,
1992).
The Liophis populations that occurred on Martinique, Marie-Galante, and St. Lucia have suffered extirpation correlated with the presence of the introduced mongoose (Herpestes auropunctatus). Liophis cursor
is now restricted to mongoose-free Rocher du Diamant and has a total range of 0.2 km2. Likewise the
distribution of L. ornatus is now restricted to mongoose-free Maria Major and has a total range of 0.1
km2. On Guadaloupe, Liophis juliae appears to have a
limited distribution on the southwestern portion of
Grande-Terre (Henderson, 1992; Henderson et al.,
1992), and the distribution of Liophis perfuscus on Barbados is restricted; Herpestes occurs on both islands,
A widely distributed Liophis population exists only on
253
mongoose-free Dominica (RWH, pers. obs.; A. Malhotra, pers. comm.).
Little is known about the natural history of West
Indian Liophis. All species are oviparous, diurnal,
ground-dwelling, active foragers. Michaud and Dixon (1989) provided information on the diets of 20
species of Liophis, but had no data regarding Antillean
taxa. The little information known about diets of West
Indian species was summarized in Schwartz and Henderson (1991). In this paper, we elaborate on what is
known about West Indian Liophis diets and make comparisons with L. melanotus from Trinidad and Tobago
and with Lesser Antillean Alsophis. We acknowledge
our small samples, but today encounters with West
Indian Liophis (except on Dominica) are infrequent.
The samples at our disposal were accumulated over
100+ years and several populations have already been
extirpated (i.e., sample sizes will never increase for
some populations). Other populations should be sampled only in very small numbers, if at all. We examined 10 Liophis cursor, 39 L. juliae, 4 L. perfuscus, and
43 L. melanotus (only from Trinidad and Tobago); no
specimens of L. ornatus were available.
We have no data regarding diet in L. perfuscus because the four specimens examined yielded no prey
remains. No snake had multiple prey items. The diet
of Liophis cursor is represented by six prey items: one
unidentified Coleoptera in a snake 570 mm SVL; two
Eleutherodactylus sp. from snakes 523-551 mm SVL;
and three Anolis roquet from snakes 388-620 mm SVL.
The diet of Liophis juliae is represented by nine prey
items: one unidentified insect in a snake 410 mm SVL;
seven Eleutherodactylus martinicensis from snakes 204450 mm SVL (two snakes with only eggs in their
stomachs were 190-382 mm SVL); and one Anolis oculatus from a snake 385 mm SVL. The diet of Liophis
melanotus is represented by: one cyprinodontid fish
from a snake 375 mm SVL; two Eleutheroductylus sp.
from snakes 237-322 mm SVL; two Gonatodes (probably vittatus) from snakes 251–267 mm SVL; four Bachia heteropus from snakes 248-292 mm SVL; and one
unidentified teiid from a snake 245 mm SVL.
The high percentage of anurans in the diet of Liophis
juliae (Fig. 1) is relatively unusual for a West Indian
colubrid. Lizards in general, and Anolis in particular,
comprise over 50% (by frequency) of the prey items
in most Antillean colubrids (Henderson and Crother,
1989). The major exception to this, for which we have
adequate data, is the small Hispaniolan endemic Darlingtonia haetiana (Henderson and Schwartz,1986;
Henderson et al., 1988). Although the high percentage of frogs is unusual for a West Indian colubrid,
this diet is typical of most Liophis (Vitt, 1983; Michaud
and Dixon, 1989). The proportions of frogs and anoles
found in L. cursor is much more typical of a West
Indian colubrid, In comparison, L. melanotus on Trinidad and Tobago preyed on fishes, frogs, and microteiids, not unusual prey for this species (Michaud and
Dixon, 1989). The absence or infrequency of Anolis in
the diet is typical of mainland snakes (Henderson and
Crother, 1989). The presence of invertebrates is also
extremely rare in the diets of West Indian colubrids
(Henderson and Crother, 1989); only one other primarily ingested invertebrate has been recovered from
a colubrid (Alsophis vudii; RWH, unpubl.).
254
NOTES
LITERATURE CITED
FIG. 1. A comparison of the diets of three species
of Liophis and Lesser Antillean (L.A.) Alsophis (four
species combined).
Alsophis is the only other colubrid genus in the
Lesser Antilles represented by more than one species;
it is superficially similar in habitus to Liophis and, like
Liophis, its members are oviparous, diurnal, grounddwelling, active foragers. They differ in size, however: cursor, juliae and melanotus have maximum SVLs
430-671 mm; Lesser Antillean Alsophis have maximum SVLs of 660–930 mm. The prey sample size for
Lesser Antillean Alsophis is 36 items from four species
(antiguae, antillensis, rijersmai, rufiventris) combined
(RWH, unpubl.): 13 Eleutheroductylus spp., 20 Anolis
spp., 2 Ameiva spp., and 1 Mus musculus (Fig. 1). The
frequency of Eleutherodactylus predation by Alsophis
is much higher in the Lesser Antilles than elsewhere
in the West Indies. Alsophis in the Lesser Antilles
exploit fewer prey genera (almost exclusively Eleutherodactylus and Anolis) than Alsophis in the Bahamas
or Greater Antilles (based on a total sample of ca. 250
prey items; RWH, unpubl.). On Cuba, for example,
A. cantherigerus exploits a minimum of 12 prey genera.
Colubrids on smaller islands (especially those not satellite to a large island; i.e., islands of the Lesser Antilles), including Alsophis and Liophis, encounter fewer
exploitable prey taxa (although not necessarily fewer
potential prey items). Narrower trophic niches are
probably determined by geographically-imposed limitations on the prey fauna, rather than innate prey
preferences.
Acknowledgments. — We thank the personnel at several institutions for the loan of specimens and for
permission to examine them for prey remains: David
Auth (Florida Museum of Natural History); Jose P.
Rosado (Museum of Comparative Zoology); Elyse J.
Beldon, W. Ronald Heyer, and George R. Zug (National Museum of Natural History); William E. Duellman, John E. Simmons, and Erik R. Wild (University
of Kansas Museum of Natural History). Kevin Lyman
was instrumental in producing Figure 1. Henderson’s
field work in the Lesser Antilles was funded by the
Institute of Museum Services Conservation Grant No.
IC-60084-86.
Dixon, J. R. 1989. A key and checklist to the neotropical snake genus Liophis with country lists and
maps. Smithson. Herpetol. Infer. Serv. (79):1-28
+ 12 maps.
———, and E. J. Michaud. 1992. Shaw’s black-headed snake (Liophis melanotus) (Serpentes: Colubridae) of northern South America. J. Herpetol. 26:
250-259.
Henderson, R. W. 1992. Consequences of predator
introductions and habitat destruction on amphibians and reptiles in the post-Columbus West Indies. Carib. J. Sci. 28:1-10.
———, and B. I. Crother. 1989. Biogeographic patterns of predation in West Indian colubrid snakes.
In C. A. Woods (ed.), Biogeography of the West
Indies: past, present, and future, pp. 479-518. Sand
Hill Crane Press, Gainesville, Florida.
———, and A. Schwartz. 1986. The diet of the Hispaniolan colubrid snake, Darlingtonia haetiana. Copeia 1986:529-531.
———, T. A. Noeske-Hallin, B. I. Crother, and A.
Schwartz. 1988. The diets of Hispaniolan colubrid snakes II. Prey species, prey size, and phylogeny. Herpetological 44:55-70.
———, J. Daudin, G. T. Haas, and T. J. McCarthy.
1992. Significant distribution records for some
amphibians and reptiles in the Lesser Antilles.
Carib. J. Sci. 28:101-103.
Michaud, E. J., and J. R. Dixon. 1989. Prey items of
20 species of the neotropical colubrid snake genus
Liophis. Herp. Rev. 20(2):39-41.
Schwartz, A., and R. W. Henderson. 1991. Amphibians and reptiles of the West Indies: descriptions,
distributions, and natural history. Univ. Florida
Press, Gainesville. xvi + 720 pp.
Vitt, L. J. 1983. Ecology of an anuran-eating guild
of terrestrial tropical snakes. Herpetological 39:5266.
Caribbean Journal of Science, Vol 29, No. 3-4, 254-255, 1993
copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
Additions to the Herpetofauna
of Honduras
JAMES R. MCCRANIE, 10770 SW 164th Street, Miami, Florida 33157.
The Mosquitia region of northeastern Honduras
contains one of the largest remaining tracts of lowland rainforest in Central America. However, a sizeable influx of campesinos into the region has taken
place in recent years, resulting in a considerable
amount of rainforest being felled, principally along
the major rivers and coastal areas. As the human population in Honduras increases, the pressures on the
Mosquitia region will certainly also increase. Much
of Mosquitia remains poorly known herpetologically
and there is an urgent need for survey work.
Through the efforts of Gustavo A. Cruz of the Universidad National Autonoma de Honduras, the herpetofauna of the Rio Platano basin, in the northern
NOTES
Mosquitia, is now fairly well known. Between 27 August-8 September 1992, I visited the previously uncollected environs of the Rio Wampu between its confluences with the rios Lagarto and Patuca. Six species
of amphibians and reptiles were collected for the first
time in Honduras and are reported below. Localities
are in the department of Olancho, except for Quebrada Waskista, which is in Gracias a Dios. Specimens
are deposited in the National Museum of Natural
History, Washington, DC, USA (USNM).
Hyalinobatrachium pulveratum (Peters): Confluence
of rios Sausa and Wampu, 100 m (USNM 321690; active at night on vegetation overhanging a small
stream). This specimen extends the known range ca.
240 airline km NNE from Finca Tepeyac, Matagalpa,
Nicaragua (Starrett and Savage, 1973). I follow RuizCarranza and Lynch (1991) for the generic placement
of this species.
Agalychnis saltator Taylor: Confluence of Quebrada
Siksatara and Rio Wampu, 95 m (USNM 321731; inactive at night on the trunk of a small tree ca. 2.0 m
above the ground). This specimen extends the known
range ca. 150 airline km NW from Eden Mine, Zelaya,
Nicaragua (Duellman, 1970).
Anotheca spinosa (Steindachner): Confluence of
Quebrada Siksatara and Rio Wampu, 95 m (USNM
321691; active at night on a tree branch ca. 2.0 m above
the ground). This locality is ca. 930 airline km ESE of
the Chiapas, Mexico localities for the species (Johnson
et al., 1977) and ca. 500 airline km N of the northernmost Costa Rican locality (Duellman, 1970). Anotheca spinosa is known from 500-1800 m in Mexico
(Duellman, 1970; Johnson, 1989) and from 300-1200
m in Costa Rica and Panama (Duellman, 1970). The
Honduran locality is the lowest recorded for the species.
Eleutherodactylus fitzingeri (O. Schmidt): Confluence
of Quebrada Siksatara and Rio Wampu, 95 m (USNM
321692-93); confluence of Quebrada Waskista and Rio
Wampu, 85 m (USNM 321694-706); confluence of rios
Aner and Wampu, 100 m (USNM 321707-13); confluence of rios Sausa and Wampu, 100 m (USNM 32171423); confluence of rios Yaunguay and Wampu, 110 m
(USNM 321724-30). Most specimens were taken at
night from low vegetation 0.5–2.0 m above the ground.
Large females were found at night on the ground
alongside streams. This species is one of “. . . the commonest lowland frogs in humid situations from Nicaragua through Costa Rica and Panama” (Savage, 1974:
291). Considering its abundance along the Rio Wampu, it is surprising that E. fitzingeri was not previously
reported from Honduras.
Norops oxylophus (Cope): Confluence of Quebrada
Waskista and Rio Wampu, 85 m (USNM 321732; sleeping at night on vegetation ca. 0.5 m above the ground
alongside a stream); confluence of Quebrada Siksatara
and Rio Wampu, 95 m (USNM 321733; sleeping at
night on vegetation ca. 1.0 m above the ground alongside a small stream). These specimens extend the
known range ca. 130 airline km NNW of the Bonanza,
Nicaraguan localities listed by Fitch and Seigel (1984).
255
These authors had postulated that N. oxylophus “. . .
probably does not occur farther north than Nicaragua” (p. 8).
Micrurus alleni K. P. Schmidt: Confluence of Quebrada Waskista and Rio Wampu, 85 m (USNM 32173536; both taken shortly after dark, one while inactive
underneath leaves on the forest floor and the other
as it was emerging from a hole in the ground). These
specimens extend the known range ca. 130 airline km
NNW from Bonanza, Nicaragua and ca. 200 airline
km W from Cabo Gracias a Dios, Nicaragua (Savage
and Vial, 1974). Villa (1984) plotted another locality
in extreme northern Nicaragua that is intermediate
between the two northern Nicaraguan localities mentioned above, but provided no documentation for the
record.
Acknowledgments. — I thank E. Munoz G. of COHDEFOR for providing collecting and exportation permits, Instrumental in planning the logistics of the
river trip was M. Espinal and J. Porras O. and J. Rindfleish provided field assistance. I especially thank E.
Flores, who provided expert service as a guide and
collector. Comparative material and assistance with
the identification of some of the specimens was provided by J. Savage.
LITERATURE CITED
Duellman, W. E. 1970. The hylid frogs of Middle
America. Monogr. Univ. Kansas Mus. Nat. Hist.
1:i-xi, 1-753.
Fitch, H. S., and R. A. Seigel. 1984. Ecological and
taxonomic notes on Nicaraguan anoles. Milwaukee Pub. Mus. Contrib. Biol. Geol. 57:1-13.
Johnson, J. D. 1989. A biogeographic analysis of the
herpetofauna of northwestern Nuclear Central
America. Milwaukee Pub. Mus. Contrib. Biol. Geol.
76:1-66.
———, C. A. Ely, and R. G. Webb. 1977. Biogeographical and taxonomic notes on some herpetozoa from the Northern Highlands of Chiapas,
Mexico. Trans. Kansas Acad. Sci. 79(3-4):131-139
(1976).
Ruiz-Carranza, P. M., and J. D. Lynch. 1991. Ranas
Centrolenidae de Colombia I. Propuesta de una
nueva clasificacion generica. Lozania 57:1-30.
Savage, J. M. 1974. On the leptodactylid frog called
Eleutherodactylus palmatus (Boulenger) and the status of Hylodes fitzingeri O. Schmidt. Herpetological
30(3):289-299.
———, and J. L. Vial. 1974. The venomous coral
snakes (genus Micrurus) of Costa Rica. Rev. Biol.
Trop. 21(2):295-349.
Starrett, P. H., and J. M. Savage. 1973. The systematic
status and distribution of Costa Rican glass-frogs,
genus Centrolenella (Family Centrolenidae), with
description of a new species. Bull. So. California
Acad. Sci. 72(2):57-78,
Villa, J. 1984. The venomous snakes of Nicaragua:
a synopsis. Milwaukee Pub. Mus. Contrib. Biol.
Geol. 59:1-41.
256
NOTES
Caribbean Journal of Science, Vol. 29, No, 3-4, 256-258, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
Description of the Tadpole of
Hyla catracha (Anura: Hylidae)
JAMES R. MCCRANIE, 10770 SW 164th Street, Miami, Florida 33157.
LARRY D AVID W ILSON , Department of Biology, MiamiDade Community College, South Campus, Miami, Florida
33176.
K ENNETH L. WILLIAMS , Department of Biology, Northwestern State University of Louisiana, Natchitoches, Louisiana 71497.
Porras and Wilson (1987) described Hyla catracha
from several cloud forest localities in southwestern
Honduras and adjacent El Salvador. Wilson and
McCranie (1993) provided a key to 43 of the 47 Honduran species of anurans with free-living larvae. Hyla
catracha was one of four species for which tadpoles
were unknown. Later, we collected several lots of
tadpoles in various stages, metamorphosing individuals, and recently transformed juveniles of H. catracha
at the type locality (USNM 316541-44; Zacate Blanco,
Departamento de Intibuca). Identical tadpoles (USNM
316545-46; 17.5 km SW Tutule, Departamento de La
Paz) also were collected at the other published Honduran locality for the species (Porras and Wilson, 1987).
We herein provide a description of the H. catracha
tadpole. Terminology follows Wilson and McCranie
(1993). Institutional abbreviations are listed in Leviton et al. (1985).
Typical tadpole (Fig. 1) in developmental stage 39
(Gosner, 1960) (measurements in mm): body length
11.6; tail length 21.0; total length 32.6; body slightly
depressed, height/width 0.78; snout rounded in dorsal aspect and in profile; eyes moderately large (eye
diameter 1.4; eye diameter/body height 0.23), separated by distance of 3.0, directed laterally; nostrils
situated at a point slightly closer to eyes than tip of
snout, directed anterolaterally; spiracle sinistral, directed posteriorly, situated slightly below midline at
a point about two-thirds of distance from tip of snout
to posterior end of body; vent tube dextral; caudal
musculature moderately robust, extending nearly to
tip of rounded tail, with height of caudal musculature
at midlength of tail slightly higher than dorsal fin,
about same height as ventral fin; dorsal fin extending
slightly onto body,
Oral disc (Fig. 2) medium-sized (oral disc diameter
3.5; oral disc diameter/snout width 0.76), directed
ventrally, completely bordered by two rows of moderately-large marginal papillae (ca. 18-20/mm); a single row of similar-sized submarginal papillae anterior
and posterior to jaw sheaths, submarginal row merging with additional similar-sized papillae lateral to
edges of tooth rows; oral disc not emarginated; keratinized jaw sheaths well developed, bearing discrete, rounded serrations; upper jaw sheath arched,
with long, slender lateral processes; lower jaw sheath
shallowly V-shaped; labial tooth rows 2/3, with second anterior row narrowly interrupted medially; anterior tooth rows subequal, long, extending to lateral
portion of oral disc; posterior tooth rows subequal,
nearly as long as anterior tooth rows.
In life, the tadpoles have brown bodies without
conspicuous markings, the caudal musculature is
creamy tan with pale brown mottling laterally, and
the tail fins are transparent with brown mottling.
Tadpoles in stages 26-39 have identical mouthparts
to the one described above, whereas tadpoles in stage
40 have a wide gap in the A-2 tooth row. By stage 41,
all tooth rows are fragmented and the lower jaw sheath
and the marginal papillae on the median portion of
the anterior labium have been lost. By stage 42, the
only remains of the tadpole oral disc parts are the
marginal papillae lateral to the former position of the
edges of the tooth rows, and on the posterior labium,
although the papillae are much reduced in size, A
typical tadpole in stage 26 (USNM 316541) has a body
length of 9.2 and a tail length of 13.4; one in stage
36 (USNM 316544) has 12.7 and 19.6, respectively, A
tadpole in stage 42 (USNM 316544) has a body length
of 10.6 and a tail length of 19.3; one in stage 43 (USNM
316543) has 9.9 and 10.6, respectively. Several tadpoles in stage 44 (USNM 316543) have body lengths
of 10.5-11.0, several in stage 45 (USNM 316543-44)
have body lengths of 10.5-10.7. A metamorphosed
individual in stage 46 (USNM 316543) has a body
length of 11.2. Color in life for specimens in stages
42-46 and slightly older individuals (USNM 316543)
is as follows: dorsal surfaces of body and limbs bright
FIG. 1. Lateral view of the tadpole of Hyla catracha (USNM 316544) in Gosner stage 39.
NOTES
257
FIG. 2. Oral disc of the tadpole of Hyla catracha (USNM 316544) in Gosner stage 39.
iridescent green in some and bronze-brown in others;
hidden parts of limbs yellow in all; venters gray with
white pustules in all; iris copper brown in all. The
older specimens in USNM 316543 were recognizable
as H. catracha and were similar in color to larger juveniles and adults collected at the same time from
bromeliads on hillsides above the stream occupied by
the tadpoles. A slightly larger juvenile (USNM 316540,
SVL 15.4) collected in a bromeliad between 2100-2150
m had the following coloration in life: dorsum dark
bronze; dorsal surfaces of limbs bronze, hidden portions yellow; venter gray with white pustules; iris
copper.
The tadpoles and recently metamorphosed individuals from the type locality were collected 16-17
August 1987 from an artificial pool in a small, slowmoving stream in a pasture at 2070 m. Many recently
metamorphosed individuals were perched on vegetation on the banks of this pool on the night of 16
August. Rana berlandieri tadpoles also were found in
the pool. A few H. catracha tadpoles, along with one
of Hypopachus barberi, were also taken from mud beneath a large rock in a nearby, formerly water-filled
depression, The tadpoles from SW of Tutule were
taken in slow-moving streams and associated pools
on 20-21 August 1987 at 2060-2070 m. Hyla catracha
was heard calling from impenetrable second-growth
vegetation along one of these streams on the night
of 20 August.
Porras and Wilson (1987) concluded that H. catracha
is closely related to members of the H. pinorum group
of Duellman (1970) and is most closely related to H.
melanomma. The tadpole of H. melanomma (see Duellman, 1970:400 and Figs. 201, 202) is very similar to
that of H. catracha. The only seemingly significant
difference between the two is that H. melanomma tadpoles have five posterior tooth rows instead of three
in H. catracha. The habitat of both tadpoles is similar,
Duellman (1970:401) reported that tadpoles of H. melanomma were found “. . . in shallow. pools in small
rivulets. ” Taylor (1940) reported that adults and newly metamorphosed juveniles of H. melanomma were
taken from bromeliads. Likewise, juvenile H. catracha
retreat to bromeliads shortly after metamorphosing
(pers. observ., also see Porras and Wilson, 1987), and
adult H. catracha are frequently encountered in bromeliads during the day. The overall morphological
similarities of the tadpoles and the similar natural
histories of H. melanomma and H. catracha suggest a
close relationship between the two species, in agreement with conclusions based on the adult morphology (Porras and Wilson, 1987).
Hyla catracha tadpoles will key to couplet 25 in the
key provided by Wilson and McCranie (1993), but not
to either Plectrohyla guatemalensis or Smilisca sordida in
that couplet. To incorporate H. catracha into the key,
we offer the following amendment to this couplet:
Papillae in submarginal row anterior and
posterior to jaw sheaths larger than those in
marginal rows; size moderately large, total
length 43-49 mm in stage 36 . . . . . . . . . .
. . . . . . . . . . . . . . . . Plectrohyla guatemalensis
Papillae in submarginal row anterior and
posterior to jaw sheaths, if present, similarsized to those in marginal rows; size moderately small, total length about 32 mm in
stage 36 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25A
25A. Serrated edge of upper jaw sheath forming
a continuous arch with lateral processes; a
distinct row of submarginal papillae anterior and posterior to jaw sheaths
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hyla catracha
25.
258
NOTES
Serrated edge of upper jaw sheath medially
notched, not forming a continuous arch with
lateral processes; no row of submarginal papillae anterior and posterior to jaw sheaths
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Smilisca sordida
Acknowledgments. — We are grateful to G. and C. Cruz
for their help in obtaining the collecting permits from
the Departamento de Recursos Naturales Renovables
and to J. Porras Orellana for many favors. KLW was
aided in his field work by a grant from Northwestern
State University.
LITERATURE CITED
Duellman, W. E. 1970. The hylid frogs of Middle
America. Monogr. Mus. Nat. Hist. Univ. Kansas
(1):1-753.
Gosner, K. L. 1960. A simplified table for staging
anuran embryos and larvae with notes on identification. Herpetological 16(3):183-190.
Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E.
Dawson. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and
ichthyology. Copeia 1985(3):802-832.
Porras, L., and L. D. Wilson. 1987. A new species of
Hyla from the highlands of Honduras and El Salvador. Copeia 1987(2):478-482.
Taylor, E. H. 1940. Herpetological miscellany No. I.
Univ. Kansas Sci. Bull. 26:489-571.
Wilson, L. D., and J. R. McCranie. 1993. Preliminary
key to the known tadpoles of anurans from Honduras. Royal Ontario Mus., Life Sciences Occas.
Pap. (40):1-12.
Caribbean Journal of Science, Vol. 29, No. 3-4, 258-261, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
Skull Morphology and Diet of
Antillean Bat Species
A RMANDO RODRIGUEZ-DURAN, Department of Biology,
Inter American University, Minillas Industrial Park, Bayamon, Puerto Rico 00959.
A LLEN R. LEWIS AND Y ARISA M ONTES , Department of
Biology, University of Puerto Rico, Mayaguez, Puerto Rico
00680.
In their discussion of the use of morphology for
describing patterns of organization within species assemblages, Miles et al. (1987) stress the importance
of verifying the association between ecology and
morphology before attribution of ecological significance to indirect morphological indices. A relationship between diet and skull morphology is generally
accepted (e.g., Fenton, 1982; Hiiemae and Crompton,
1985). The relationship is statistically significant for
pools of species of bats from diverse areas (Freeman,
1979, 1981, 1984, 1988). Freeman proposed the hypothesis that insectivorous species with more robust
skulls eat harder insects. However, under some circumstances skull morphology may bear more directly
on what an organism is capable of eating than on
what it actually eats. The realized diet depends on
food availability, which is affected by abundance and
behavior of both prey and competing consumers.
Given the relatively depauperate insect faunas of
islands and the possible ecological release experienced by insular populations of widespread vertebrates (Hespenheide, 1973), natural assemblages of
insular vertebrates present good subjects to test the
strength of Freeman’s ideas. The guild of insectivorous bats in Puerto Rico was used to test the hypothesis. We established statistically the position of each
species relative to others in terms of skull robustness
and looked at how well skull robustness predicted
the hardness of prey consumed by each bat species.
We also assessed intraspecific variation and sexual
dimorphism in skull morphology.
The species analyzed are sympatric in the subtropical moist-forest (Ewel and Whitmore, 1973) of the
karst belt in northwestern Puerto Rico. We included
all the insect-eating bats known to occur in the area:
Eptesicus fuscus, Pteronotus quadridens, P. parnellii, Mormoops blainvilli, Tadarida brasillensis, Molossus molossus,
and Monophyllus redmani. The latter species was included because results from a previous study demonstrated that insects are important in its diet (Rodriguez-Duran and Lewis, 1987).
We prepared six adult skulls for each species except
E. fuscus, of which we obtained only one individual.
All individuals used in this study were adults captured in north and western Puerto Rico, between the
towns of San German and Quebradillas. Preparation
of the skulls followed Silva-Taboada (1979). We took
14 measurements on each specimen using a dial caliper and a dissecting microscope, or with a camera
lucida following Freeman (1981). These measurements were converted to ratios to eliminate the effects
of size (Freeman, 1981; Sokal, 1965). Measurements
taken were: maxillary tooth row (MTR) over condylocanine length, height of upper canine over MTR, anterior-posterior thickness of upper canines over MTR,
maxillary width at the upper canines over greatest
width at the upper third molars, length of M3 over
MTR, length of the premetacrista of M3 over MTR,
length of the post-paracrista of M 3 over MTR, length
of the cusp row of P4-M 3 over MTR, area of P4-M 3
over MTR2, length of dentary condyle to protoconid
of M1 over dentary length (DL), width of M 3 talonid
over width of M3 trigonid, lateral dentary thickness
at the protoconid of M2 over DL, height of coronoid
process over DL, and height of dentary condyle above
the lower toothrow over DL. Mean values of skull
measurements taken for each species are provided in
Table 1.
We used the principal components analysis from
the SPSS program (Nie et al., 1975) to analyze these
fourteen measurements and obtain factor scores for
each bat species (Kleinbaum and Kupper, 1978). A
food hardness index, obtained by assigning a hardness value to each insect order (Freeman, 1981), was
regressed against these scores (Sokal and Rohlf, 1981).
For bats present on continents and on the islands,
only data on diet from the islands was used to obtain
NOTES
259
TABLE 1. Average value of skull measurements (mm) taken for each species. Sample sizes consist of six
individuals of each species, except for E. fuscus which is represented by one female. M.m. = M. molossus, T.b.
= T. brasiliensis, M.r. = M. redmani, E.f. = E. fusscus, M.b. = M. blainvillii, P.q. = P. quadridens, P.p. = P. parnellii.
Maxillary toothrow
Condylocanine
Upper canine
Maxillary width
Width at M3
Thickness canine
Length M3
Premetacrista M3
Postparacrista M3
Cusprow
Area cusprow
Dent. cond-protocon.
Dentary length
M 3 talonid
M 3 trigonid
Protoconid M3
Coronoid proc.
Dentary condyle
M.m.
T.b.
M.r.
E.f.
M.b.
P.q.
P.p.
6.4
15.7
2.3
4.5
8.2
1.6
0.9
—
0.7
4.7
1505
8.7
11.96
0.6
0.9
1.2
3.9
1.8
5.7
14.2
1.7
4.2
6.9
1.2
0.95
0.7
0.7
3.9
1055
7.6
10.4
0.8
0.8
0.8
3.3
1.8
6.8
17.6
1.8
3.3
4.7
1.3
0.8
0.3
0.4
3.5
377
7.7
13.2
0.5
0.5
0.6
3.2
1.96
7.2
16.7
2.6
5.9
8.1
1.7
0.97
0.5
0.6
5.1
1386
10.1
14.5
0.7
0.97
1.2
5.9
3.2
7.7
13.1
2.7
4.5
6.4
1.5
0.7
0.7
0.66
4.1
1108
6.5
12.3
0.7
0.8
0.6
3.4
3.2
6.0
13.1
1.9
3.9
5.5
1.1
0.8
0.6
0.55
3.6
827
6.3
10.1
0.6
0.7
0.5
2.6
2.3
8.6
18.1
2.5
4.8
7.4
1.7
1.1
0.8
0.67
5.4
1748
9.3
15.0
0.8
0.9
0.9
3.8
3.5
blainvillii 49 SCS, 65 FS; P. parnellii 47 SCS; M. redmani
65 SCS, 64 FS; E. fuscus 25 SCS, 3FS; T. brasillensis 27
SCS; M. molossus 26 SCS, 41FS. Since what an individual bat eats must vary both temporally and spatially, the way we assessed diet is more reliable than
matching each skull with the stomach content of the
individual from which the skull was obtained. Data
on diet for each species are provided in Table 2.
Factor 1 of the PC accounted for 56.5% of the variance in the data and separated the bats into a group
the food-hardness score. Information on diet was obtained from Silva-Taboada (1979) (stomach content
samples, SCS) Rodriguez-Duran and Lewis (1987) and
this study (fecal samples, FS). Information on diet was
obtained from both the rainy and dry seasons. Fecal
samples obtained for this study, as well as those reported by Rodriguez-Duran and Lewis (1987), were
from the same area where the bats were collected.
The number of samples used to obtain food-hardness
score are as follows: P. quadridens 195 SCS, 158 FS; M.
T ABLE 2. Diet of insectivorous bats used to determine’ food hardness score. Numbers in the columns
represent the percentage of fecal pellets containing that item. Sources of information are labeled as: 1, This
study (Fecal samples); 2. Rodriguez-Duran and Lewis, 1987 (Fecal samples); and 3. Silva-Taboada, 1979 (Stomach
samples). Sample sizes are provided in the text. PP = Pteronotus parnellii, PQ = P. quadridens, MB = Mormoops
blainvillii, EF = Eptesicus fuscus, TB = Tadarida brasiliensis; MM = Molossus molossus, and MR = Monophyllus
redmani.
Insect order
Lepidoptera
Orthoptera
Diptera
Coleoptera
Odonata
Hymenopteran
Dictyoptera
Hemiptera
Homoptera
Dermaptera
Neuroptera
Ephemeroptera
PP
3
93.0
14.3
21.4
14.3
14.3
7.1
PQ
MB
EF
2
3
2
3
84
17.6
16.2
41.2
75.0
90
100.0
14
45
5.3
45
83
16
47
35
1.0
1.0
1.0
13.2
1.0
1
100
3
1
3
12
25.9
58.5
19.2
38.4
12
72
70.7
7.4
7.4
29.6
7.3
82.9
8
19
15
100
MM
TB
3
8
10.5
62
24
34
9.8
48.9
18.5
11.5
MR
2
26.9
11.5
65.4
3.8
4
7.6
4
10
260
NOTES
FIG. 1. Regression of skull robustness (factor score
on PC1) on food hardness. The regression line is based
on the average factor score for each species and is not
significantly different from zero with a one-tailed test
(P = 0.05). In this figure Y = 2.83 + 0.45X, t = 1.891,
squares represent males, crosses represent females.
composed of M. molossus and E. fuscus, and a second
group of five species (Fig. 1). Low intraspecific variation in skull morphology was demonstrated by the
tight clusters of individuals of each species along PC1.
The species with most intraspecific variation (M. molosSUS) also showed the strongest trend toward sexual
dimorphism. The variables that loaded heavily on
factor 1 are similar to those found by Freeman (1981)
to be most important in defining factor 1 in her analysis. They were: ratio of talonid/trigonid of M 3
(–0.8745), relative height of upper canine (0.8573),
relative anterior-posterior thickness of upper canine
(0.8422), relative thickness of dentary (0.7847), relative height of coronoid process (0.6669), relative length
of the premetacrista of M3 (–0.6499), and relative
maxillary width at the upper canines (–0.6030). We
used factor 1 as the index of skull robustness, thus
allowing comparison with Freeman’s results.
The slope of the regression of food-hardness on
skull robustness, based on the average robustness value for each species (Fig. 1), was analyzed with a onetailed test. Based on Freeman’s work, species with
relatively robust skulls consume relatively hard prey.
If Freeman’s result applies to Puerto Rico, the relationship between skull robustness and food hardness
should be positive. The observed t value (t = 1.891,
df = 5, P = 0.059) does not permit rejection of the
null hypothesis that the slope is less than or equal to
zero. However, the probability of the observed t is
small enough that the results do not justify rejecting
Freeman’s result for insectivorous bats in Puerto Rico.
This conclusion results in part from the observation
that with only seven species of insectivorous bats in
western Puerto Rico, a very strong relationship would
be necessary for a statistically significant regression.
It is also prompted by the observation that the coefficient of determination of the regression (R 2 = 0.42)
is similar to the value obtained by Freeman (1981) (R2
= 0.38). Coefficients of determination are unbiased
by sample size (McNeil et al., 1975; Edwards, 1984).
One species in particular, P. quadridens, probably
accounts for the failure to outright accept Freeman’s
hypothesis. The species scored low on skull robustness but fed on hard foods: predominantly coleopterans (Silva-Taboada, 1979; Rodriguez-Duran and
Lewis, 1987) and hymenopterans.
A discrepancy between the skull robustness of some
bats and the hardness of their food, suggests that
morphological trends reveal only part of the answer
to the question of what influences the diet of bats.
Freeman (1981) cites other examples where there is
discrepancy between what bats are capable of eating
and what their morphology suggests they should eat.
Time, foraging strategy, space, and social behavior
may influence diet (Fenton, 1982). Nevertheless, morphology is a powerful indicator of both function and
adaptive value, and useful structural hypotheses can
be derived from morphological data (Webster and
Webster, 1988). Our data does not point toward a
looser association between skull morphology and diet
for island-dwelling bats than for continental species.
Further study based on large assemblages, rather than
pools of species from different geographical areas, is
warranted.
Acknowledgments. — We thank P. W. Freeman for
clarification of the measurements taken and two
anonymous reviewers for their comments. This work
was partly funded by ASM Sigma Xi (Boston U. Chapter), Inter American University of Puerto Rico Grantsin-Aid of research to AR-D and the NIH-MBRS Program through NIAMS Grant No. S06RR08103 to ARL.
The University of Puerto Rico, Mayaguez Campus
provided Computer time, and the Department of Natural Resources of Puerto Rico provided permits to
capture bats. Myrna Vega kindly typed the manuscript. Gladys Otero from the office of the Associate
Dean for Research, Mayaguez, prepared the figure.
LITERATURE CITED
Edwards, A. L. 1984. Multiple regression and the
analysis of variance and covariance. Freeman, New
York. 221 pp.
Ewel, J. J., and J. L. Whitmore. 1973. The ecological
life zones of Puerto Rico and the U.S. Virgin Islands. Forest Serv. Research Pap. ITF-8, USDA, 72
pp. + map.
Fenton, M. B. 1982. Echolocation, insect hearing,
and feeding ecology of insectivorous bats. In T. H.
Kunz (ed.), Ecology of bats, pp. 261-286. Plenum
Press, New York.
Freeman, P. W. 1979. Specialized insectivore: beetleeating and moth-eating molossid bats. J. Mamm.
60:467-479.
———. 1981. Correspondence of food habits and
morphology in insectivorous bats. J. Mamm. 62:
164-166.
———. 1984. Functional cranial analysis of large
animalivorous bats (Microchiroptera). Biol. J. Linn.
Soc. 21:387-408.
———. 1988. Frugivorous and animalivorous bats
(Microchiroptera): dental and cranial adpatations.
Biol. J. Linn. Soc. 33:249-272.
Hespenheide, H. A. 1973. Ecological inferences from
NOTES
morphological data. Ann. Rev. Ecol. Syst. 4:213229.
Hiiemae, K. M., and A. W. Crompton. 1985. Mastication, food transport, and swallowing. In M. Hildebrand et al. (eds.), Functional vertebrate morphology, pp. 262-290. Harvard University Press,
Cambridge, Massachusetts.
Kleinbaum, D. G., and L. L. Kupper. 1978. Applied
regression analysis and other multivariable methods. Duxbury Press, Boston, Massachusetts. 556
pp.
McNeil, K. A., F. J. Kelly, and T. T. McNeil. 1975.
Testing research hypotheses using multiple linear
regression. Southern Illinois University Press,
Carbondale, Illinois. 587 pp.
Miles, D. B., R. E. Ricklefs, and J. Travis. 1987. Concordance of ecomorphological relationships in
three assemblages of passerine birds. Amer. Nat.
129:347-364.
Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner,
and D. H. Bent. 1975. Statistical package for the
social sciences. McGraw-Hill, New York. 675 pp.
Rodriguez-Duran, A., and A. R. Lewis. 1987. Coexistence in a narrow-mouthed cave: patterns of
population size, diet, and activity time for a multispecies assemblage of bats. Carib. J. Sci. 23:352360.
Silva-Taboada, G. 1979. Los murcielagos de Cuba.
Editorial Academia, La Habana, Cuba. 429 pp.
Sokal, R. R. 1965. Statistical methods in systematic.
Biol. Rev. 40:337-391.
———, and F. J. Rohlf. 1981. Biometry. W. H. Freeman & Co., New York. 859 pp.
Webster, D. B., and M. Webster. 1988. Hypotheses
derived from morphological data: when and how
they are useful. Amer. Zool. 28:231-236.
Caribbean Journal of Science, Vol. 29, No. 3-4, 261-262, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
Territorial Aggression by a
Migrant Tennessee Warbler:
Defense of an Artificial
Food Source
ROBERT L. NORTON , 961 Clopper Road, Apt. B1, Gaithersburg, Maryland 20878.
Migrant landbirds experience annual variations in
resources during migration and upon reaching breeding and wintering areas. Occasionally, a migrant may
be late or off-course from its migratory route. Survival
during abnormal conditions requires a flexible strategy to accommodate novel, unusual, potentially unfavorable environmental conditions (Zumeta and
Holmes, 1978) and competition from residents and
migrants experiencing similar constraints (Woolfenden, 1962; Kale, 1967). Plasticity of behavior, and prey
or resource choice among migrating warblers has been
discussed (Wunderle, 1978; Greenberg, 1986; Scaly,
261
1988; Morse, 1989) as well as causes for fluctuating
resource availability (Scaly, 1988). Short-term territoriality may be beneficial to an individual facing the
survival threshhold (Schemske, 1975). However, Emlen (1973) proposed that territorial defense of nectar
resources by a migrant insectivore would be maladaptive. Greenberg (1986) states that territoriality is
common, even among immature warblers, and may
bean adaptive strategy among several groups of birds.
Observations of wood warbler behavior during migration (Kale, 1967; Trainer and Trainer, 1977; Wunderle, 1978; Scaly, 1988) focus on the behavioral strategy of securing a resource, or territory, by employing
aggressive displays, or chases, generally towards conspecifics. In this note I describe aggressive behavior
of a wood warbler during spring migration at an unnatural resource—a feeder.
On 17 March 1986 at Cruz Bay, St. John, U.S. Virgin
Islands, I observed during an approximately 30 min
period a Tennessee Warbler (Vermivora peregrina) vigorously defending a sugar-feeder used primarily by
Bananaquits (Coereba flaveola). There were frequent
chases by V. peregrina, and repeated striking (1-3 jabs
with its beak) of the cranium of bananaquits which
crowded at the feeder. Intense physical aggression is
not generally noted of Tennessee Warblers during
migration, but is apparently evident in wintering areas (Stiles, 1983).
The “territorialism” during migration that I observed at an artificially rich resource was apparently
invoked as a “switched-on” or “fine-tuned” behavior
strategy in response to the quality of the resource
(Leek, 1972; Morton, 1979). The stimulus for this behavior (social intolerance) was the result of niche
similarity (i.e., commensal with a nectar source), and
possibly hierarchal dominance (Morse, 1974), or both
(Greenberg, 1986).
Environmental conditions on St. John played a role
in a conditioned aggression response, as it apparently
does in wintering areas. The climate during the first
quarter of 1986 in the Virgin Islands was very dry
(about 45% below average rain fall [Norton, 1987]).
Near the Hobbs’ feeder station in Cruz Bay, aloe (Aloe
vera) and century plants (Agave americana) are typically in full bloom during March and provide a rich
source of nectar and insects for birds (Emlen, 1973;
pers. obs.). In the Virgin Islands, wood warblers are
usually found in high concentrations in moist forests
(Askins et al., 1992; Ewert and Askins, 1991; Norton,
1979, 1981) and mangrove forests (Robertson, 1962).
Since a normally rich source of nectar was lacking in
March 1986, and an artificial resource was being exploited by a gregarious, resident nectivore, defense
of a resource-rich feeder by this species is not unexpected.
Tennessee Warblers have shown a persistent, or
commensal, relationship with Combretum fructcosum
blossoms (Trainer and Kemp, 1979; Morton, 1982) and
territorial behavior when resources are reduced during winter (Morton, 1982), yet somewhat gregarious
behavior in intraspecific feeding groups when resources are abundant (Morton, 1980). Moynihan (1962)
and Leek (1972) suggested that migrants are generally
joiners in resident intraspecific feeding flocks in the
tropics, but are frequently the recipients of aggressive
262
NOTES
behavior (Willis, 1966; Rappole et al., 1983) when food
resources are limited. Vrwivora peregina apparently
invokes aggression, or social intolerance, during migration in contrast to joiner behavior in winter.
Acknowledgments. — I thank Ann and Joe Hobbs
for their consideration and hospitality; David Spector,
Eugene Morton, and Russell Greenberg for preliminary review and suggestions; Joseph Wunderle and
Kevin Winker for critical review.
LITERATURE CITED
Askins, R. A., D. Ewert, and R. L. Norton. 1992.
Abundance of wintering warblers in fragmented
and continuous forests in the Virgin Islands. In J.
A. Hagan III and D. Johnston (eds.), Ecology and
conservation of neotropical migrant landbirds, pp.
197-206. Smith. Inst. Press, Washington, DC.
Emlen, J. T. 1973. Territorial aggression in wintering
warblers at Bahama Agave blossoms. Wilson Bull.
85:71-74.
Ewert, D., and R. A. Askins. 1991. Flocking behavior
of migratory warblers in winter in the Virgin Islands. Condor 93:864-868.
Greenberg, R. 1986. Competition in migrant birds
in the nonbreeding season. In R F. Johnston (ed.),
Current ornithology, Vol. 3. pp 281-307. Plenum
Press, New York.
Kale II, H. W. 1967. Aggressive behavior by a migrating Cape May Warbler. Auk 84:120-121.
Leek, C. F. 1972. Seasonal changes in feeding pressures of fruit and nectar eating birds in Panama.
Condor 74:54-60.
Morse, D. H. 1974. Niche breadth as a function of
social dominance. Am. Nat. 108:818-830.
———. 1989. American warblers an ecological and
behavioral perspective. Harvard Univ. Press,
Cambridge, Massachusetts. 406 pp.
Morton, E. S. 1979. Effective pollination of Erythrina
fusca by the orchard oriole (Icterus spurius): coevolved behavioral manipulation? Ann. Missouri
Bot. Gard. 66:482-489.
———. 1980. Adaptations to seasonal changes by
migrant landbirds in the Panama Canal Zone. In
A. Keast and E. S. Morton (eds.), Migrant birds in
the Neotropics: ecology, behavior, distribution, and
conservation, pp. 437-457. Smith. Inst. Press,
Washigton, DC.
———. 1982. Adaptations for tropical survival in
wintering North American birds, Nat. Geog. Soc.
Research Report. Vol. 14:491-498.
Moynihan, M. 1962. The organization and probable
evolution of some mixed species flocks of neotropical birds. Smith. Misc. Coll. 143(7):1-140.
Norton, R. L. 1979. New records of birds for the
Virgin Islands. Amer. Birds 33145-146.
———. 1981. Additional records and notes of birds
in the Virgin Islands. Amer. Birds 35:144-147.
———. 1987. The spring migration, West Indies region. Amer. Birds 40:528-529.
Rappole, J. H., E. S. Morton, T. E. Lovejoy, III, and J.
L. Rous. 1983. Nearctic avian migrants in the
neotropics. U.S. Dept. of Int., Fish and Wildl. Serv.,
Washington, DC. 646 pp.
Robertson, W. B. 1962. Observations on the birds of
St. Job., Virgin Islands. Auk 79:44-76.
Schemske, D. W. 1975. Territoriality in a nectar feeding Northern Oriole in Costa Rica. Auk 92:594595.
Scaly, S. G. 1988. Aggressiveness in migrating Cape
May warblers defense of an aquatic food source.
Condor 90:271-274.
Stiles, G. F. 1983. Vermivora peregrina. In D. H. Janzen
(ed.), Costa Rican natural history, pp. 613-614.
Univ. Chicago Press, Chicago.
Tramer, E. J., and T. R. Kemp. 1979. Diet-correlated
variations in social behavior of wintering Tennesse Warblers. Auk 96:186-187.
———, and F. E. Tramer. 1977. Feeding responses
of fall migrants to prolonged inclement weather.
Wilson Bull. 89:166-167.
Willis, E. O. 1966. The role of migrant birds at ant
swarms. Living Bird 5:187-231.
Woolfeneden, G. E. 1962. Aggressive behavior by a
wintering Myrtle Warbler. Auk 79:713-714.
Wunderle, J. M., Jr. 1978. Territorial defense of a
nectar source by a Palm Warbler. Wilson Bull. 90:
297-299.
Zumeta, D. C., and R. T. Holmes. 1978, Habitat shift
and roadside mortality of Scarlet Tanagers during
a cold wet New England spring. Wilson Bull. 90:
575-586.
Notes on Breeding of the
Puerto Rican Tanager
(Nesospingus speculiferus)
RAUL A. PEREZ-RIVERA, Department of Biology, University of Puerto Rico, Humacao, Puerto Rico 00792.
Little is known about the biology of the endemic
Puerto Rican Tanager (Nesospingus speculiferus). The
scant information on its breeding biology is anecdotal
and is summarized by Isler and Isler (1987). Gundlach
(1882) reported a nest found by A. Stahl attributed to
the Puerto Rican Tanager. The nest was described as
a cup constructed of grass and feathers, and lined with
thin grass. This is the same nest described by Danforth
(1936), Bond (1980), and Biaggi (1983). The latter author added that it contained a bluish-white egg (26
× 18.5 mm) with brown speckles, and a few black
spots and streaks. Raffaele (1989) reported for the species a cup-shaped nest and a clutch of 2–3 creamcolored eggs heavily speckled with dark brow. Here
I provide a detailed description of nests and eggs of
the Puerto Rican Tanager. I also present data on nesting sites and on the breeding season of the species
within the Carite Forest.
The Carite Forest is located in southeastern Puerto
Rico (Fig. 1). It is classified as a subtropical moist forest
with an annual rainfall of 100-200 cm and temperatures between 18 and 24°C (Ewell and Whitmore, 1973).
263
NOTES
FIG. 1. Location of the study area (shaded portion).
The study area was located at an altitude of 750 m.
The forest at this location is a two-strata formation.
The dominant vegetation includes candlewood (Dacryodes excelsa), sierra palm (Prestoea montana), wild
mango (Micropholis chrysophylloides) and dovewood
(Alchornea latifolia).
Since 1975, I have made incidental observations on
the breeding biology of the Puerto Rican Tanager.
During this period, I have found four nests. Location
and measurements of nests are presented in Table 1.
Nest A was under construction on 10 January 1981
in a 3-m Sloanea berteriana shrub, The nest was being
constructed from vines, roots of epiphytes, and fibers
of sierra palm leaves. The nest-tree was cut down five
days later, and the incomplete structure was destroyed. Nest B was discovered on 11 September 1983
by the calls of a pair of tanagers with three fledglings.
The nest was in a 3-m Mecranium amigdalinum shrub.
It was an open cup with soft, yet substantial walls. It
was formed with the same material as the previous
one and included fungal hyphae of Marasmius spp.
The nest was lined with fibers of the leaves of the
sierra palm and some downy tanager feathers. The
suspended cup was sewn with fungal hyphae to the
vines at the rim. Nest C (Fig. 2) was found on 21
March 1992 in an 11-m high dovewood tree. It was
placed near the distal end of a branch covered with
vines of Ipomoea (Convolvulaceae). The nest was in
the final stage of construction. Both adults were observed carrying nesting material, although the female
did most of the building. This nest was constructed
of the same materials as the previous ones, but also
included parts of leaves and stems of the ferns Microgram piloselloides and Grammitis spp. It was lined
exclusively with thin strips of sierra palm leaves. This
nest was also sewn with fungal hyphae to the vines
TABLE 1. Location and measurements of Puerto Rican Tanager nests at Carite Forest.
Nests
Measurements
1
Location of nesting tree
Location of nest
Height of nest above ground
B
C
D
BF
DEB3
2.0
BF
DEB
2.5
BF
DEB
10.0
BF
DEB
3.0
—
—
—
—
7.0
4.5
9.0
7.5
6.5
5.0
10.0
7.8
6.0
4.5
8.5
7.0
A
2
Nests
Outside height
Depth of cup
Outside diameter
Diameter of cup
1
Measurements in cm, except for height of nest (m).
Border of forest (BF).
3
Distal end of branch covered with vines (DEB).
2
264
NOTES
with my data. The breeding season of this tanager at
Carite, coincides with those of most other forest birds
at this and other interim localities in Puerto Rico
(Perez-Rivera, 1986).
Acknowledgments. — I thank Efrain Nadal and Gilbert Bonilla for assisting in the field work. Eugenio
Santiago, Vicente Quevedo and Gary Breckon identified some of the nest materials. Eli Espinosa is responsible for Fig. 2. I am particularly grateful to Wayne
J. Arendt and Elizabeth Hodges for improving preliminary drafts of the manuscript and to unknown
reviewers for their helpful suggestions.
LITERATURE CITED
FIG. 2. Nest of a Puerto Rican Tanager (Nesospingus
speculiferus).
at the rim. Nest C contained three whitish subelliptical eggs boldly scribbled or scrawled and with reddish to dark brown blotches and spots The eggs measured 23.1 × 17.1 nun, 23.8 × 17.3 mm, and 24.1 ×
17.4 mm. Nest C, as well as one egg, is in the collection
of the Museum of the University of Puerto Rico in
Humacao. Nest D was found on 25 April 1992 in a
3.5-m dovewood shrub. The structure was similar in
location and construction to nest C. It was abandoned
before egg laying.
All four nests were cup-shaped as the ones described by Gundlach (1882) and Raffaele (1989), although none were constructed with grass. Roots of
epiphytes, vines, fungal hyphae and strips from the
leaves of the sierra palm (used as lining) were a common component of all nests studied. These materials
are not mentioned in previous works.
The eggs that I examined do not match the description in Raffaele (1989) nor the one (including the
dimensions) of Stahl (described in Gundlach, 1882;
Biaggi, 1983). Either the eggs of the Puerto Rican
Tanager range in color like those of the Blue-gray
Tanager (Thraupis episcopus) (Isler and Isler, 1987), or
were confused in previous studies with those of the
Stripe-headed Tanager (Spidalis zena portoricensis). The
latter species constructs variable bowl k. cup-shaped
nests and lays 2-3 eggs that vary in color and markings (Perez-Rivera, 1992).
At Carite, I have found active nests of the Puerto
Rican Tanager in January, March, April and September. I have observed tanagers stripping leaves of the
sierra palm for nest construction in April and May at
El Yunque and Maricao, respectively. The tanagers,
at least in Carite, defend nesting territories usually
from late December to the end of July. From late July
to mid-August, most Puerto Rican Tanagers abandon
their nesting territories and join foraging flocks. In
the Carite Forest, the courtship song of the species is
most commonly beard from March to June, and is
infrequent after July. The tanagers at Carite probably
nest at least twice a year.
Raffaele (1989) reports the Puerto Rican Tanager
breeding from January to August, which is consistent
Biaggi, V. 1983. Las Aves de Puerto Rico, 3rd ed.
Editorial Universitaria, Universidad de Puerto Rico,
Rio Piedras, Puerto Rico. 373 pp.
Bond, J. 1980. Birds of the West Indies. Houghton
Mifflin Co., Great Britain. 210 pp.
Danforth, S. T. 1936. Los Pajaros de Puerto Rico.
Rand McNally and Co., New York. 168 pp.
Ewell, J., and J. Whitmore. 1973. The ecological life
zones of Puerto Rico and the U.S. Virgin Islands.
U.S.D.A. Forestry Service Res. Paper ITF-18. Institute of Tropical Forestry, Rio Piedras, Puerto
Rico. 72 pp.
Gundlach, J. 1882. Briefliches: zur Fortpflanzungs
geschichte des Chlorospingus speculiferus. J. Ornithol. 30:161.
Isler, M. L., and P. R. Isler. 1987. The Tanagers:
natural history, distribution and identification.
Smithsonian Institution Press, Washington, D.C.
404 pp.
Perez-Rivera, R. A. 1986. Parasitism by the Shiny
Cowbird in the interior parts of Puerto Rico. J.
Field Ornithol. 57:99-104.
———. 1992. Variabilidad en los nidos y huevos de
la Reina Mora. El Pitirre. 5(3):14.
Raffaele, H. A. 1989. A Guide to the Birds of Puerto
Rico and the Virgin Islands. Princeton University
Press, Oxford, United Kingdom. 254 pp.
Carribean and of Science, Vol. 29, No. 3-4, 264-267, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
Preservation of a
Clypeasteroid Echinoid in
Holocene Beachrock, Jamaica
STEPHEN K. DONOVAN , ROGER A. WILLIAMS , AND JUDY
A. ROCKE , Department of Geology, University of the West
Indies, Mona, Kingston 7, Jamaica.
Beachrock is produced predominantly in tropical,
carbonate-rich, intertidal environments by penecontemporaneous cementation in the zone between high
and low tides (Scoffin, 1987). Beachrock is commonly
composed of mainly carbonate grains with a carbonate (aragonite and/or high-magnesium calcite; Stoddart and Cann, 1965; Meyers, 1987) cement, but non-
NOTES
265
FIG. 1, A. Map of The Palisades and adjacent area, southern Jamaica. Key: A = Norman Manley lnternational Airport; HR = Hope River; K = Kingston; KH = Kingston Harbor; PR = Port Royal. Localities 1
and 2 are arrowed. Locality 1 (GR 786435) is the site where Clypeaster rosaceus was collected by J.A.R. as a
reworked cobble of beachrock. Photographs in Fig. 2 were taken at locality 2 (GR 725420). B. Outline map of
Jamaica showing position of study area (arrowed). Key: K = Kingston; O = Ocho Rios; M = Montego Bay.
carbonate beaches can similarly be lithified (Scoffin,
1987). We report herein the occurrence of an unusual
beachrock lithology enclosing an unexpected bioclast.
The specimen described here was collected from
The Palisadoes, parish of St. Andrew, Jamaica (Fig.
1), a tombolo (Hendry, 1979) which forms the southern boundary of Kingston Harbor and occurs to the
northwest of the Yallahs Basin (Burke, 1967). Hendry
(1979) noted that in situ beachrock occurs almost everywhere on the seaward shoreline of The Palisades.
Beach and beachrock sediment in The Palisades is
polymict, varying from sands to boulders. Individual
beds within beachrock are up to 0.5 m thick (Fig. 2A),
with a seaward dip of 2° to 6° and a micritic cement
of magnesian calcite (Hendry, 1979). In contrast, the
beachrock formed in the limestone environment of
the nearby Port Royal Cays (Fig. 1) is cemented by
aragonite (Hendry, 1979). Beachrock illustrated in Fig.
2 outcrops at locality 2, while the echinoid specimen
described below was collected from beach rubble at
locality 1 (Fig. 1).
Of significance is the occurrence in The Palisades
beachrock, which varies between a sandstone and a
boulder conglomerate in lithology, of well-lithified
marine invertebrate skeletons. The presence of manade artifacts indicates that lithification of at least
the upper beds was very recent (certainly 20th century); hence, the lithified invertebrate skeletons enclosed therein are therefore of similar age. The most
abundant skeletons were those of more massive scleractinian corals (Fig. 2B), large gastropod (Fig. 2C, D),
oysters and other benthic molluscs. Hendry (1979)
recorded the occurrence of rare, thin horizons of unbroken mollusc shells.
A test of the extant clypeasteroid echinoid Clypeaster rosaceus (Linne) has been collected as a cobble
from the beach at locality 1 (Fig. 1), but is obviously
derived from the beachrock (Fig. 3). This species is
locally common in the Pleistocene of Jamaica and has
been reported from a number of localities (Donovan,
1993). This species also lives at least close to the study
area, around the Port Royal Cays (S.K.D., pers. obs.),
although snorkel observations have not been made
close to The Palisadoes. Most significantly for the
present study, it is known (from the August Town
Formation in the Hope River Gorge (Chubb, 1958;
Donovan and Lewis, 1993) to the northeast of the
study area (Fig. 1). Perhaps a test of Clypeaster could
be reworked out of the August Town Formation to
be washed up on the south coast of The Palisadoes
after fluvial and nearshore transport. However, the
lithology of the sediments in filling the test show it
to have been derived from the local beachrock. Despite the well-know” strength of the test of Clypeaster
(Donovan, 1991), it is questionable whether it would
have been so well-preserved after several kilometres
of transport.
The test is abraded, so that details of the apical
surface are indistinct, but the five petals, the apical
266
NOTES
FIG. 2. Lithoclasts and bioclasts preserved in exposed beachrock at locality 2 (Fig. 1A), The Palisadoes,
Jamaica. A. Exposed section through part of the beachrock succession; note alternation of sandstone and
conglomerate horizons. B-D. Bioclasts incorporated into conglomeratic horizons. B. Platey scleractinian coral.
C. Gastropod Cassis sp. (arrowed). D. Gastropod Strombus sp. (arrowed). The hammer is 280 mm in length.
system and some primary tubercles are preserved.
The ambitus has been abraded away, exposing the
internal infill of a terrigenous, polymict sedimentary
rock with gravel-sized clasts in a matrix/cement of
mainly carbonate. This rock also obscures the oral
surface. Internal supports of the test are visible in the
ambital region (Fig. 3).
Scoffin (1970) recorded a conglomeratic beachrock
from the Bahamas in which the clasts comprised shells
of Strombus, bottles, fragments of coral and boulders
of limestone with carbonate sand. In contrast, the
polymict beachrock at The Palisadoes is comprised of
sand- to boulder-sized lithoclasts derived by the erosion, fluvial and longshore transport of a variety of
igneous, sedimentary and metamorphic lithologies
from the Blue Mountain massif to the east, as well as
locally-derived bioclasts with some man-made artifacts The lithoclasts are transported to the coast by
the Hope (Fig. 1A) and Yallahs Rivers (Wescott and
Ethridge, 1980), with longshore drift occurring in
an east to west direction (Hendry, 1983). Scoffin (1987)
noted that beachrock lithification of non-carbonate
sands occurs on tropical volcanic islands, but The
Palisadoes deposit is unusual for the large range of
lithologies shown by the incorporated clasts. This is
a function of the diversity of rock types in the source
area, although at least some of the boulders may have
been derived from ships, ballast.
The presence of boulder-sized clasts makes the
preservation of an echinoid test in this deposit all the
more unexpected. We know of no other report of an
echinoid preserved in such an environment, although complete echinoids have been reported from
other high-energy deposits. Kidwell and Baumiller
(1990) and Greenstein (1991) demonstrated the resilience of at least some regular echinoid tests during
tumbling transport, so perhaps the preservation of a
robust test such as that of C. rosaceus in a high-energy
267
NOTES
F I G. 3. Clypeaster rosaceus (Linne), The Natural
History Museum, London, EE3000, from Holocene
(20th century) beachrock. The Palisadoes, parish of
St. Andrew, Jamaica (locality 1 in Fig. 1). Lateral view
(anterior to left) to show abraded ambital region and
infill of lithic fragments with a carbonate cement.
Arrow indicates internal support of test. ×0.7.
deposit should not be unexpected. However, despite
the presence of other tax. washed up on the beach
as dead tests (including Echinometra lucunter (Linne)
and E. viridis A. Agassiz, regular echinoids with particularly robust tests; Gordon 1991; Greenstein, 1991),
no other echinoids were observed in the beachrock,
despite a modest abundance of other benthic taxa
such as molluscs and corals.
Other Holocene beachrocks have also preserved
unusual bioclasts (Easton, 1974). The significance of
The Palisadoes occurrence is to indicate the potential
for fossil echinoderms to be well-preserved in ancient
beachrock horizons. As beachrock has been identified
as far back as the Proterozoic (Donaldson and Ricketts,
1979), there is a real possibility that ancient beachrocks may preserve complete skeletons of robust,
shallow sub-littoral echinoderms that were washed
up onto the beach and rapidly lithified.
Acknowledgments. — We thank Hal Dixon for his help
in the field and George Soloman for photographing
the echinoid specimen. This research was undertaken
during the period of a Shell Distinguished Research
Fellowship in Science to S.K.D., which is gratefully
acknowledged. We thank David Lewis and an anonymous referee for their constructive comments.
LITERATURE CITED
Broke, K. 1967. The Yallahs Basin: a sedimetary
basin southeast of Kingston, Jamaica. Mar. Geol.
5:45-60.
Chubb, L. J. 1958. Higher Miocene rocks of south
east Jamaica. Geonotes 1:26-31.
Donaldson, J. A., and B. R. Ricketts. 1979. Beachrock
in Proterozoic dolostone of the Belcher Islands
Northwest Territories, Canada. J. Sedim. Petrol
49:1287-1294.
Donovan, S. K. 1991. The taphonomy of echinoderms: calcareous multi-element skeleton in the
marine environment. In S. K. Donovan (ed.), The
processes of fossiliization. pp. 241-269 Belhaven
Press, London.
———. 1993. Jamaican Cenozoic Echinoidea. In R.
M. Wright and E. Robinson (eds.), Biostratigraphy
of Jamaica. Geol. Soc. Am. Mem. 182. In press.
———, and D. N. Lewis. 1993. The H.L. Hawkins
collection of Caribbean fossil echinoids annotated
catalog of rediscovered specimens from the University of Reading, England. Carib. J. Sci. 29:186201.
Easton, W. H. 1974. An unusual inclusion in beachrock. J. Sedim. Petrol. 44:693-694.
Gordon, C. M. 1991. The poor fossil record of Echinometra (Echinodermata: Echinoidea) in the Caribbean region. J. Geol. Soc. Jamaica 28:37-41.
Greenstein, B. J. 1991. An itegrated study of echinoid taphonomy: predictions for the fossil record
of four echinoid families. Palaios 6:519-540.
Hendry, M. D. 1979. A study of coastline evolution
and sedimentology The Palisadoes, Jamaica. Ph.D.
Thesis. University of the West Indies, Mona. 232
pp.
———. 1983. The influence of the sea-land breeze
regime on beach erosion and accretion: an example from Jamaica. Carib. Geogr. 1:13-23.
Kidwell, S. M., and T. Baumiller. 1990. Experimental
disintegration of regular echinoids: roles of temperature, oxygen, and decay thresholds. Paleobiol.
16:247-271.
Meyers, J. H. 1987. Marine vadose beachrock cementation by cryptomystalline magnesian calcite—Maui, Hawaii. J. Sedim. Petrol. 57:558-570.
Scoffin, T. P. 1970. A conglomeratic beachrock in
Bimini, Bahamas. J. Sedim. Petrol. 40:756-759.
———. 1987. An introduction to carbonate sediments and rocks. Blackie, Glasgow. 274 pp.
Stoddart, D. R., and J. R. Cann. 1965. Nature and
origin of beach rock. J. Sedim. Petrol. 35:243-247.
Wescott, W. A., and F. G. Ethridge. 1980. Fan-delta
sedimentology and tectonic setting—Yallahs fan
delta, southeast Jamaica. Am. Ass. Petrol. Geol.
Bull. 64:374-399.
Caribbean Journal of Science, Vol. 29, No. 3-4, 267-271, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
Key for Field Identification of
Rudist Bivalves from the
Cretaceus of Jamaica
ELIZABETH R. DAVIS -STRICKLAND , Palm Beach Community College, 3000 Saint Lucie Avenue, Boca Raton, Florida
33431-6490.
STEPHEN K. DONOVAN , Deportment of Geology, University of the West Indies, Mona, Kingston 7, Jamaica.
Field biostratigraphy is usually limited by the proficiency of the investigator, so many stratigraphically
useful or even important fossils may be overlooked
due to ignorance. This is unfortunate, as any fossil
may give useful data pertaining to the age of the
enclosing rocks if only it can be released by the investigator. This normally requires large collections of
fossils to be made for later identification in the lab-
.
TABLE 1. Key to the morphology and taxonomy of the fossil rudist bivalves of the Cretaceus of Jamaica
(based on Chubb, 1971; with some supplementary data from Perkins, 1969; McFarlane, 1977; authors’ personal
observations). Key: E = encruster; J = juvenile; X = clinger; a = Biradiolites sp. cf. B. rudissmus Trechmann is
known from the Campanian; b = vascular markings; c = many curved shells may also be straight (in some
species curvature is limited to the juvenile stages only); d = “. . . diameter, dorso-ventral excluding the flange
33 mm, including flange 62 mm, antero-posterior 47 mm . . .“ (Chubb, 1971:190); e = discoidal or saucershaped; f = “Right [=attached] valve subcylindrical, slightly curved or straight. . .“ (Chubb, 1971:204); g =
right (attached) valve not known or not described from Jamaica; h = internal mould of right valve only; i =
T ABLE 1. Extended.
external surface of Monopleura sp. unknown; j = external sculpture of varix-like annulations; k = a vesicular
l
tissue lines the body cavity, rather than regular tabulae; = Biradiolites and Bournonia have “quadrangular
prismatic cells” (Dechaseaux and Coogan, 1969:N810) in right valve, but Chubb only reported this from
Biradiolites forbesi amongst the Jamaican species; m = recorded in one individual only (Chubb, 1971:200),
otherwise absent; n = irregular, rounded tetragonal and highly lobate in outline; closed circle = character
present; ? = character possibly present; empty space = character absent.
NOTES
270
oratory. Making such a collection has limitations, such
as the damage done to the collecting site, the time it
takes to make a representative collection and weight.
The last problem is particularly important in the Upper Cretaceus of the Caribbean region, where the
dominant macrofossils (numerically and in biomass)
in shelf carbonate sequences are often the unusual
bivalve molluscs (superfamily Hippuritacea) commonly known as rudists. That rudists are frequently
giants among benthic molluscs is well-known. Typical Caribbean genera include Titanosarcollites Trechmann, which grows to over 2 m in total length, and
Bournonia Fischer, that can exceed 0.6 m in diameter
(Perkins, 1969:N751). Such specimens are not easy to
collect.
In an attempt to produce a simple field identification chart to the Jamaican rudist fauna, we have been
inspired by the key to the morphology and taxonomy
of the bivalves published by Skelton (1985a:Table
6.4.1). This enables identification of any bivalve to
the level of superfamily. Our tabulation (Table 1) enables any of the rudists found in Jamaica to be identified to species, by reference to relatively few morphological features of the shell. While limited to taxa
reported from Jamaica by Chubb (1971), this table is
broadly applicable to the Caribbean.
EXPLANATIONS
Table 1 considers morphological attributes of rudist
species recorded from Jamaica by Chubb (1971). The
features recorded are, as far as possible, those actually
known from each species. Only in a few cases have
diagnostic generic characteristics not recorded from
Jamaican species been included; these may be misleading because some of the species are in need of
generic revision (P. W. Skelton, pers. comm.). The
characters examined in Table 1 are those generally
easiest to see. These characters are largely self-explanatory, but the accounts of Perkins (1969), Chubb
(1971) and, particularly, Donovan (1992) are recommended for additional definitions. The following
points are considered worthy of further explanation:
(1) Rare. — Specimens described by Chubb (1971)
from five or less specimens, or which are simply called
‘rare’ in Chubb’s monograph.
(2-5) Size. — The diameter at the commissure of
complete adult shells (where known) are used
throughout (Chubb, 1971:165). Rudists are rarely circular, so the dorso-ventral and anterio-posterior diameters are usually different. The size ranges indicated are adult maxima. See Chubb (1971) for
discussions of shell heights.
(25) Attached. — Herein includes clingers (= ’encrusters’; Skelton, 1985b) and elevators sensu Skelton
(1991).
(26) Recumbent. — Recumbent is used sensu Skelton
(1985b, 1991) and includes large, free-living, unattached forms which enclose a large virtual area.
(27-33) External features. — Includes features from
the right, left or both valves, as available.
(27) Flange. — Projections more or less perpendicular to the shell surface. Titanosarcolites alatus Chubb
has flanges at least 46 mm in length (Chubb, 1971:
178).
(28) Grooves. — A narrow furrow on the shell surface.
(29) Striations. — Fine grooves on the surface, giving
the shell a striated appearance.
(30) Costae/Ridges. — A rib on the shell surface,
somewhat less pronounced than a flange.
(31) Smooth. — A shell lacking external sculpture
(with the possible exception of fine growth lines).
(32) Infoldings. — The folding inwards of the outer
shell surface.
(33) Growth lines. — These occur on the thin (often
not preserved) outer shell layer.
(34-41) Internal features. — Features of the shell,
rather than of the mantle cavity. Chubb (1971) did
not describe or figure the internal features of all the
Jamaican species. Terms such as tubes, tubules, etc.,
were used interchangeably, at least to some extent.
(34) Oscules. — Tubes within the free valve which
channeled water to the mantle cavity (for a discussion of function, see Skelton, 1976).
(35) Accessory cavity. — Cavity between the myophores (sites for the attachment of adductor muscles)
and the inner shell wall.
(36) Tubes. — Usually hollow (occasionally septate)
structures within the shell wall, teeth or hinge area.
(37) Tubules. — Small tubes.
(38) Septa. — Plates or dividing platforms in the
tubes, tubules or canals.
(39) Polygonal canals. — System of canals of four or
more sides formed by anastomosizing vertical plates
in the shell wall.
(40) Prismatic cells. — Cells formed between the funnel plates and the vertical walls.
(41) Capillaries. — Very fine tubules.
(42-45) Mantle cavity. — Chubb (1971) did not describe or figure the body cavity of all of the species
considered herein.
(44) Reniform. — Kidney-shaped.
(45) Tabulae. — The bottom of the body cavity is
floored by a septum. Tabulae is used rather than septate to avoid confusion with (38), above.
The following points may be of importance when
used in association with Table 1. Some specimens of
Bournonia and Thyrastylon show no septa in the body
cavity due to diagenetic replacement. ‘Vascular markings’ are found on the commissure in Birdiolites forbesi, B. rudissimus and Durania nicholasi. Chiapasella radiolitiformis, Praeradiolites coatesi and Durania nicholasi
may have vesicular structures in the body cavity. Plagioptychus and Mitrocaprina have a brown exterior. Mitrocaprina multicanaliculata, Plagioptychus toucasianus and
Sphaerucaprina seafieldensis are described from the free
valve only. Barrettia is distinguished from Parastroma
by a ‘string of beads’ (=outer shell infoldings) apparent on the commissure, although this may not be
apparent in all species.
LITERATURE CITED
Chubb, L. J. 1971. Rudists of Jamaica. Palaeontogr.
Am. 7(45):157-257.
Dechaseaux, C., and A. H. Coogan. 1969. Family
Radiolitidae Gray, 1848. In R. C. Moore (ed.), Treatise on invertebrate paleontology, part N, Mollusca 6(2), pp. N803-N817. Geological Society of
NOTES
America and University of Kansas Press, Boulder
and Lawrence.
Donovan, S. K. 1992. A plain man’s guide to rudist
bivalves. J. Geol. Ed. 40:313-320.
McFarlane, N. 1977. Jamaica 1:250,000 geological
sheet. Ministry of Mining and Natural Resources,
Kingston.
Perkins, B. F. 1969. Rudist morphology. In R. C.
Moore (ed.), Treatise on invertebrate paleontology, part N, Mollusca 6(2), pp. N751-N764. Geological Society of America and University of Kansas Press, Boulder and Lawrence.
Skelton, P. W. 1976. Functional morphology of the
Hippuritidae. Lethaia 9:83-100.
271
———. 1985a. Class Bivalvia. In. J. W. Murray (ed.),
Atlas of invertebrate macrofossils, pp. 81-101.
Longman, Harlow, England.
———. 1985b. Preadaptation and evolutionary innovation in rudist bivalves. Spec. Pap. Paleont. 33:
159-173.
———. 1991. Morphogenetic versus environmental
cues for adaptive radiations. In N. Schmidt-Kittler
and K. Vogel (eds.), Constructional morphology
and evolution, pp. 375–388. Springer-Verlag, Berlin.
BOOK REVIEWS
Caribbean Journal of Science, Vol. 29, No. 3-4, 272, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
Threatened Birds of the Americas. N. J. Collar,
L. P. Gonzaga, N. Krabbe, A. Madrono Nieto,
L. G. Naranjo, T. A. Parker and D. C. Wege.
1992. Smithsonian Institution Press, Washington, D.C. 1150 paginas, varios mapas y
dibujos. ISBN 1-56098-267-5. Carpeta dura $75.
Este libro es una estupenda contribucion a la conservacion de la avifauna de las Americas. Es una excelente guia que traza prioridades de conservacion y
recoge recomendaciones para el manejo y preservacion de especies de aves y sus habitats amenazados o
en peligro de extincion. Sera de mucha utilidad para
oficiales de gobierno, organizaciones de conservacion, estudiantes y especialistas en aves, todos con
una responsabilidad o mandato que exige un uso responsible y racional del patrimonio natural mundial.
Este libro es el segundo en una serie proyectada de
cuatro volumenes que forman parte de la tercera edicion del Red Data Book preparado por el Consejo
International para la Preservacion de Aves (I.C.B.P.,
siglas en ingles). La meta fundamental de los autores
fue buscar, analizar e incluir todo material relevante
a la conservacion de las especies de aves amenazadas
o en peligro de extincion en las Americas, y organizarlo en un informe de conservacion. Considerando
lo extenso de la lista de especies, el esfuerzo de los
autores por hacer acopio de la informacion y presentarla en forma organizada, fue sencillamente monumental.
La presentacion sigue un formato estandard. Aparece el nombre comun de la especie (en negritas) y
el nombre cientifico, seguido de una sinopsis de la
informacion presentada (en bastardillas). Siguen secciones que tratan los temas de la distribucion, poblaciones, ecologia, amenazas a la supervivencia, medidas de proteccion, conservacion y comentarios. La
seccion de distribucion detalla la extencion geografica
de las especies. La seccion de poblaciones discute el
status, abundancia y tendencias poblacionales. La seccion de ecologia cubre aspectos del habitat, alimentos
y reproduccion. En la seccion de amenazas a la supervivencia se identifican amenazas potenciales y actuales que ponen en riesgo la preservacion de las
especies. La seccion de medidas para la proteccion y
conservacion tiene recomendaciones concretas sobre
el manejo y proteccion de las especies y sus habitats.
En esta seccion se identifican otras especies simpatridas con la especie bajo consideracion. Hay una seccion
de comentarios que discute conflictos en la interpretacion y analisis de la informacion, presenta testimonios sobre informacion adicional y provee comentarios taxonomicos sobre las especies.
Resulta muy practico presentar listas de especies,
agrupadas en categorias de status (de paso fijando
prioridades para su conservacion) y en regiones geo-
graficas (paises) como se hace en los apendices B y C.
El apendice D presenta una lista de especies que aparentemente no estan en serio peligro de extincion,
pero que su situacion es preocupante. El libro termina
con una extensa y muy util lista de referencias, seguida de un no menos util indice de especies.
Un detalle tecnico muy importante es el sistema de
prioridades para la conservacion de las especies basado en el grado o riesgo de extincion. Para esto se
establecio un sistema numerico paralelo al tradicional
sistema de categorias de status usado en la serie de
Red Data Books.
Sobresale que este informe de conservacion fue
traducido al espanol, gracias al auspicio de la Comision para la Celebracion del Quinto Centenario (gobierno Espanol). Esta es una magnifica aportacion para
la divulgacion de este tratado tecnico entre los paises
hispanoparlantes.
Lamentablemente no se pudieron incluir en el libro
las especies amenazadas o en peligro de extincion en
America del Norte (E.U.A. y Canada) e islas del “Pacifico Neotropical.” Para subsanar esta deficiencia se
incluye en el apendice A un resumen de la situacion
de 25 especies de las regiones antes mencionadas.
Aunque no se proveen mapas de distribucion para
todas las especies incluidas, si se proveen para especies con un patron de distribucion particularmente
interesante o fuera de lo usual. No hay una lista de
mapas adjunta al cuadro de contenido. Los mapas no
contienen informacion sobre longitud y latitud en sus
margenes ni ofrecen referencia a escalas. La informacion sobre localidades (longitud y latitud) si esta
contenida en el texto de la especie bajo consideracion.
Este libro debe formar parte de la biblioteca del
ornitologo professional y de profesionales en areas
relacionadas a la conservacion.
CARLOS A. DELANNOY , Departamento de Biologia, Universidad de Puerto Rico, Mayaguez, Puerto Rico 00681
Caribbean Journal of Science, Vol. 29, No. 3-4, 272-274, 1993
Copyright 1993 College of Arts and Sciences
University of Puerto Rico, Mayaguez
272
Herpetofauna Mexicana. Lista Anotada de las
Especies de Anfibios y Reptiles de Mexico,
Cambios Taxonomicos Recientes, y Nuevas
Especies. Annotated List of the Species of
Amphibians and Reptiles of Mexico, Recent
Taxonomic Changes, and New Species. Oscar Flores-Villela (edited by C. J. McCoy).
1993. Special Publication 17, Carnegie Museum of Natural History, Pittsburg. iv + 73
pp. ISBN 0-911239-42-1 (paper). $15.00.
Seventeen years have passed since the last species
list of Mexican amphibians and reptiles appeared.
During that time new species have been described
BOOK REVIEWS
from Mexico, taxonomic and nomenclatural changes
have occurred, and several previously described species have been reported from Mexico for the first time.
Thus, a current species list of the Mexican herpetofauna is a welcome addition to the literature.
The Flores-Villela publication (abbreviated FV) is
bilingual (Spanish and English) and in parallel text,
which will facilitate its use by Spanish speaking and
English speaking people interested in the herpetofauna of Mexico. The work is divided into two parts,
a current species list with broad geographical distribution categories and a summation of recent taxonomic changes.
The introduction to the list begins with a brief discussion of the history of herpetology in Mexico and
an overview of Mexican geography and climatic zones.
FV then divides the country into 10 natural regions
by simplifying the system prepared by West in 1971
(based primarily upon climate and vegetation), and
then uses these natural regions to categorize the herpetofauna geographically. FV then describes the geographical composition of each of his 10 natural regions.
Some of these descriptions are detailed and agree
largely with his accompanying map (Regions 2, 3, 5,
7, 10), others include erroneous statements (Region
1: western Durango is not in this region; Region 6:
central Chiapas is not in this region; Region 8: eastern
Durango should read western Durango; Region 9:
north-northwestern San Luis Potosi [English text only]
is not in this region), while omissions occur in other
descriptions (Region 1: also includes most of Zacatecas and the northern one-half of Aguascalientes;
Region 4: though stated in the text that the Sierra
Madre del Sur extends into Michoacan, the map does
not indicate this). Also, the figure legend for the FV
map should read West, 1971b, not 1970b.
Following the natural regions discussion is the species list. FV’S Table 1 summarizes the number of taxa
by families, genera, and species within each order or
suborder. Several discrepancies exist between the figures in Table 1 and the actual list (195 species of
anurans given in Table 1 versus 197 listed, 337 species
of lizards versus 339, 85 genera of snakes versus 87,
and 40 species of turtles versus 39). I scrutinized the
amphibian and snake sections of the list, as these are
the groups I am most familiar with. The following
comments pertain only to these two groups, unless
otherwise stated.
I found considerable faults in the use of parentheses around authors names following the valid species
names. Linnaeus, 1758 should not be in parentheses
following Boa constrictor, whereas the authors of the
following names should have been placed in parentheses: Bufo marinus, Eleutherodactylus angustidigitorum,
E. dennisi, E. dilatus, E. grandis, E. interorbitalis, E. leprus,
E. modestus, E. nivicolimae, E. pallidus, E. pipilans, E. rubrimaculutus, E. rufescens, E. saxatilis, E. syristes, E. teretistes, E. verrucipes, Dryadophis melanolomus, Leptodeira
frenata, L. septentrionalis, Crotalus viridis and Porthidium
hespere. Erroneous dates are given for the year of authorship of the following species names (excluding
those in which the actual year of publication is uncertain or in dispute; correct year appears in parentheses): Hyla arenicolor (1866); Eleutherordactylus pallidus
(1958); Rana berlandieri (1859); Geophis isthmicus (1894);
273
Imantodes gemmistratus (1861); Salvadora lemniscata
(1895). The use of single-i or double-ii endings for
eponyms originally proposed with a double-ii is inconsistent. Thus one finds Hyla smithii, Rana boylii,
Ensatina eschscholtzii, among the amphibians, whereas
the single-i ending is adopted for Bufo luetkeni, B.
woodhousei, Agalychnis moreleti, Pseudacris clarki, Smilisca baudini, Eleutherodactylus berkenbuschi, Scaphiopus
couchi, S. hammondi, Ambystoma dumerli, Pseudoeurycea
belli, and P. gadovi. Particularly inconsistent is the use
of deppii (Cnemidophorus) or deppei (Abronia, Tantilla).
In 1992, Campbell and Smith transferred Hyla erythromma to the genus Ptychohyla and elevated P. macrotympanum to the specific level (formerly considered
a subspecies of P. euthysanota). Although FV accepted
the changes proposed by Campbell and Smith (see p.
65-66), he failed to include P. macrotympanum in his
valid species list and continued to use Hyla erythromma. In 1989, Savage and Crother placed Pliocercus andrewsi and P. bicolor in the synonymy of Urotheca elapoides, While FV may be correct in not following the
generic arrangement proposed by Savage and Crother, he states on page 68, while discussing the Savage
and Crother paper, that “. . . only the former [elapoides] occurs in Mexico.” That he includes both P.
andrewsi and P. bicolor in his species list is puzzling.
Although FV stated that his taxonomic changes are
summarized through September 1992 (p. 2), several
papers published early that year were apparently not
available to him in time to incorporate the changes
to his species list.
As stated earlier, FV’s natural regions system is a
simplified version of that offered by West (compare
Fig. 2 in FV with Fig. 3 in West’s 1971 paper). This is
particularly evident in FV’s areas 1 and 6 (1 and 11
respectively, in West’s paper) where climate, and especially vegetation, change considerably within these
respective regions to form distinctive subdivisions
(see discussion in West). There are numerous examples of amphibians and reptiles that occur only in one
or two of West’s subdivisions within either region 1
or 6 rather than being widely distributed in these
respective regions. Such a finer refinement of FV’s
natural regions would have created more work on his
part in categorizing the affected species geographic
distributions, but would have significantly improved
the usefulness of this section. Scrutiny of the natural
regions distributions of the amphibian and snake species revealed several mistakes (Ixalotriton niger should
be region 5; Crotalus triseriatus is not in region 9 if C.
aquilus is considered a distinct species) or obvious
omissions (Bufo woodhousei, region 8; Pternohyla fodiens, 7; Hypopachus variolosus, 3 [southwestern Aguascalientes]; Rana montezumae, 1 [northern Aguascalientes]; R. yavapaiensis, 2; Coluber constrictor, 8 [old record
for southwestern Durango]; Hypsiglena torquata, 8;
Lampropeltis mexicana, 8; Leptodeira annulata, 10; Masticophis taeniatus, 8[questioned by FV]; Pituophis melanoleucus, 7; Rhadinaea laureata, 8; Salvadora grahamiae,
8; Thamnophis sirtalis, 8; Trimorphodon tau, 1 [northcentral Aguascalientes]; Micrurus diastema, 3 if M. bernadi is a subspecies of M. diastema [see p. 63]; Crotalus
aquilus, 3; Crotalus pricei, 9 and 3 [northwestern Aguascalientes]; Crotalus scutulatus, 3).
The recent taxonomic changes part consists of four
274
BOOK REVIEWS
sections, viz.: species described or first recorded from
Mexico since 1966 (10+ p.); taxonomic and nomenclatural changes at the species level since 1976 (7 p.);
changes above the species level (4+ p.); changes in
type locality (1 p.). Each section contains a list of the
affected species with citations to the pertinent literature. I found these sections to be well-done and informative, as well as offering easy access to the pertinent literature concerning recent taxonomic changes
in the composition of the Mexican herpetofauna.
Most of my criticisms of this publication are relatively minor. My main concern is that the numerous
inconsistencies and the relatively few errors and
omissions in the species list will be perpetrated by
workers with limited access to the original literature.
A case in point is that the erroneous dates of authorship given by FV for Hyla arenicolor and Rana berlandieri were perpetrated from Frost’s recent list of amphibian species of the world. However, I recommend
this publication for anyone with an interest in the
herpetology of Mexico. Used in conjunction with the
works of Smith and Smith, one can access virtually
all of the literature pertaining to Mexico for any species of amphibian and reptile.
JAMES R. MCCRANIE, 10770 SW 164th Street, Miami, Florida 33157.
ANNOUNCEMENT
1994 JOINT A NNUAL M EETINGS OF THE S OCIETY FOR C ONSERVATION
B IOLOGY AND THE A SSOCIATION FOR T ROPICAL B IOLOGY
The Society for Conservation Biology and the Association for Tropical Biology will be holding their annual
meetings jointly from 7 to 12 June 1994, at the University of Guadalajara, in the state of Jalisco, Mexico. The
meeting will promote the participation of Latin American biologists in both societies and will provide a good
opportunity for “networking” and learning about conservation biology in the Neotropics. Travel, lodging
and registration costs will be similar to those of meetings held previously in the U.S.A. Information on
symposia, contributed papers, poster sessions and travel arrangements will be mailed to members of both
societies in November 1993. Deadline for submission of papers will be in early March. For additional information write or send FAX to:
Eduardo Santana or Stanley Temple
SCB-ATB Organizing Committee
Department of Wildlife Ecology
University of Wisconsin-Madison
Madison, WI 53706 U.S.A.
FAX (608) 262-6099
Bruce Benz or Enrique Jardel
SCB-ATB Organizing Committee
Laboratorio Natural Las Joyas
Universidad de Guadalajara
Apdo. Postal 1-3933
Guadalajara, Jalisco
C.P. 44100 Mexico
FAX (52-338) 7-27-49