SUCCESSION OF MAM MhALIAN FAUNAS ON
TRINIDAD, WEST INDIES
ELIZABETH KiHWARZ WING
4 Caliifi fF T.97=**NPL:iF TilL T': THE GRADUATE COUNCIL OF
ihf LIN~I.EF:IT.' OF FLORIDA
INu PF71i.11. F=.lLIL~Lr N: 5.[ HE REQUIREMENTS FOR THE
E*Fa.RF **F~ fllOCTI: I OF PHILOSOPHY
LiNriV'i.MITY' OF FLORIDA
This study could not have been done without the cooperation
of many persons and organizations to whom I am deeply indebted. I
should like to thank my graduate supervisory committee, Drs. Archie
F. Carr, James R. Redmond, and John M. Goggin, Mr. Clayton E. Ray, and
particularly my chairman, Dr. James N. Layne, for their advice and en-
couragement throughout this study. I am also most grateful for the
loan of specimens from Miss Barbara Lawrence of the Museum of Com-
parative Zoology, Dr. Richard Van Gelder of the American Museum of
Natural History, Drs. H. G. Kugler and S. Schaub of the Basel Natural
History Museum, and Mr. J. A. Bullbrook of the Royal Victoria Institute.
Field work during the summer of 1959 was carried out with the assistance
of a National Science Foundation Summer Fellowship for Graduate Teach-
ing Assistants and with the help and hospitality of Texaco Trinidad, Inc.,
and Trinidad Petroleum Development. I would also like to extend my
thanks to Dr. H. G. Kugler, Dr. K. Barr, Mr. J. A. Bullbrook, Dr. C. C.
Wilson, and Dr. I. Rouse for help in connection with various phases
of the study. Finally, I wish to express my gratitude to my parents
and husband for their help and constant encouragement.
TABLE (F CONTENTS
A.CKI~C~~DLEDGMENTS.~ .. .. . .. .. . . ..
LIST O~F TABILES . .. .. . . . . . . . . iv
LI~ST OFP ILLUTRTE~IFII ION... ..... ...-
ITFcFERODUCTION. C~X..,,.. ................ ..
P03;T-PLESTOCE202 KI;;TORY OF~ THE G;ECLGIC, CLIA4TIC, A~ND
VBGE;TAcTIiCIIAL FEATURES OF P~RUINIDJI)........
Vegetation. .. .. . . - - - - - 1i
THE PRE-BUIMI IV4UJ LAN FAUUAi iOF TRINIDAL.. . . L5
rooi3Jl MaSmmals of Prinidad. .. . . . . . . 25
~oogeographic jlignificknee of Pre-humasn FauC1a . . . ?
~MAMMAiLL PUnlA AS REPFJ'SDITE'D IN~ 17(DU.N MTfIDDiS .. .. .. j!
Description of 3ites. . .. . . . - - - 33
Intersite Comparisons .. . . - . . - 47
KISTHU.I C 1-UMIituLAN FAUNA .. .. .. .. .. . . 5-
List of' Terrestrial Idlr~ubtls .. .. . . - - 54
Zoogeograph~ic AffinitLes .. .. .. .. . .. t5
DI3CU 35ION .. . . . . . . . . . . . .
3UMMARY surI cac~CLUCS~c . . .. . . .. .. 65
LTERNRHUEf~ CITED . .. .. . . . . - - - - 73
BICI;RAPHICA~L SKEPICtl. .. . -. -.. - - - - 6
LIST OF TABLES
1 Rainfall data for selected stations on Trinidad .. 16
2 Measurements of the alveolar cheek tooth row of
various species of Zygodontom~ys . ... .. ... 29
3 Measurements of fossil _Zygodontomys .. .. ... 3C
4 Correlation of periods of occupation on Trinidad
and ceramic styles. .. .. .. .. .. .. .. .. 34
5 Relative abundance of mammals at different time
periods within the St. John site. .. ... 36
6 Relative abundance of mammals recorded from nine
middene ... .. . ... . . * 46
7 Measurements of Sipodon ef. hirsutus. .. .. .. 62
LIC.T O~F LLLUrl7PRAION3
1 Ehoreline of Trinidad, approxi~mately 17,82
y'ear s go .. . . .. .. .. .. . .. .. B
?! ,Choreline of Trinidad, approx~imately 8,iOO
years jgo .. ... .. .. ... .. .. . . 11
3 nystorgrahic regious of Trinidad .. .. .. .. 13
4 Pleistscene vegetation of~ Venezuela Fsnd Trinidsd .. 14
: Pre-Columbian vegetation of Trinijdad . .. .. .. 21
Animal remains excavated from Indian middens constitute a source
of valuable information to both the archeologist and the zoologist
(Taylor, 1957). These materials are particularly significant, since
they often can be accurately dated, in that they may provide informa-
tion about the transition between Pleistocene and Recent time, a period
in which world biotas underwent rapid changes (Burt, 1961) From the
animals represented in a midden, much about the faunal composition of
an area at a given time may be revealed. Changes in the range of cer-
tain forms may also be indicated. Such changes may in turn throw light
on the nature of past ecological conditions in a given region. Midden
remains occasionally include species that are new to science. More
often, comparison of specimens from middens with those of the same
species now inhabiting the region shows certain differences, analysis
of which may lead to a better understanding of evolutionary processes.
From an archeological standpoint, the study of animal remains from
middens contributes to a more nearly complete understanding of the
environmental milieu of the particular society under consideration.
In addition it provides data on which to base cultural hypotheses as
to the nature of hunting and butchering techniques, food taboos, and the
non-alimentary uses of animals. By integrating data on fossil and re-
cent faunas of the region a. greater perspective on this type of as-
semblage is gained and thereby a better understanding of' it.
Because of itsi dynmwic geological nister"; and attendant climatic
and vegetaitional changes, Trinidad, West Indies, is of considerable zo-
oarcheological interest. It nee become repaErated fro~m the usinlandP of'
fourtn Ame~rica, frocm whicn its faun8 and aboriginnl population were de-
rived iince it beccame inhabited by man.
'The present stuldy i- an attempt to apply the zooarcheological
approach in an analys~is of the mammalian remains from~ abo~riginal sites
in Trinidad. rThese ;ites encOmp~ass a t iae-span froml app~roximately 2751j
yearc: ago to historic times In addition, an effort has been made to
int~erate Unis faulnal assemblag~e urlta data On both yre-bu~aR and his-
toric crammials in order to gain Esoe knowledge of the hietary of the
msmmalian faulna of PriniJad from t~he PleistoceneF to tne present day as
related to g~eologic, ecologic, and human factors.
MATERIALS AN~D METHODS
The materials upon which this study is based come from a
number of sources. Many of the fossil finds on Trinidad have been
made by Dr. H. G. Kugler in the course of his career as oil geologist
for Texaco Trinidad, Inc. In 1922, he discovered a vertebrate-fossil-
bearing stratum of oil sands while making a test pit at Apex (Trinidad)
Oilfields, Inc. near Fyzabad. The fossils that were collected are in
the Basel Natural History Museum collections. A similar deposit near-
by at the Forest Reserve of Texaco Trinidad, Inc. was found in 1957
when the site was being cleared for oil well Number 1060. An almost
complete skeleton of Glyptodon was excavated, and shipped along with
some fragments to the American Mu~seum of Natural History. None of
this material has yet been prepared or catalogued. A carbon-14 date
based upon wood associated with Megatherium bones was also taken at
My husband and I conducted field work in Trinidad during the
summer of 1959 vith the objectives of collecting more fossil, midden,
and Recent vertebrate material. and of getting first hand knowledge of
the ecology of the island. All specimens collected by us are deposited
in the appropriate University of Florida Collection. An attempt to re-
locate the fossil locality at Apex was unsuccessful in spite of help
from the Apex Company. At the site at Forest Reserve, several bones
of larger mammals were collected, and a stratum containing small ver-
tebrate fossils was discovered. The small fossils were extracted from
the oil sands in which they were embedded by soaking large clumps of it
fcr Jevcrail days in gaaoinre, thren brilinrg the clump8 to bre~ak them
up, and finally 'rnshing the residue? tnrough creensJ. Initialil 20~,
160j, and3 230 gauge screensj were us~ed but since very few fo~ssls wenrt
through thje 20 Gauge screen, subsequent b~atches were washed only
through a 5il gaurEc screen.
Toe bulk: of the midden mlaterials wai excavated by Dr. 1. Rouse,
Yale University,, snd Dr. J. H. G~oZgin, University~ of Florida, in 13k(
and 1953, under the auspices of' the Ylale Caribbean Research Program
and Graduate schooll of the University of Florida. even sites--!.
Josepb, Maylo, -t. John, Cedros, Elri, Palo Seco, and Quinaml--weer
represented in these collections. Phe excavations were made be re-
mocving one or more sections twoc meters square. Each Fection was
dividedj vertically into a number of levels 20 cm. in depth. The bones
as well as the artifacts from teac level were kept separate, so that
stratigrapnic changes cou~ld subsClequentl be analyzedj. Another col-
lection of midden bones which dBE BVailabiD for studjy irns made by
Dr. Kagler at Cedros. Phis was not an excavaticon, but rattner a 5ur-
face collectionr. Additioal~ masterial came from an excavation made at the
Erin site by Mlr. J. A. bullIOrook, of the Royal. Victoria Institut~e. ;l-
though this site had been excavated in a manner similar to that employed
by Rouse and G~oggin, the bones vere not kept segreg~ated intc- their
In tte cauim3r of 1959, I visited adx known archeological rites
in Trinidad for the purposes of making ecological observations at
each sits and to collect additional surficial material. The sites
studied included Chagonaray, Palo Seco, Mayaro, Guayaguayare, Mayo,
and St. John.
The zooarcheological material obtained from Trinidad middens
was in quite good condition compared to similar material from
other areas. Only about 50 percent of the material was too frag-
mentary to permit identification. Of the identifiable component,
about 50 percent was mammal bones, and the remainder was fish, rep-
tile, and bird. The mammal elements identified totaled 4884. In this
study the'hbinimum number of individuals" is used as an index of the
relative abundance of a particular species in a sample of midden
remains. For each species this is determined by the count of the
most numerous skeletal element in a given lot. In some cases this
number may validly be increased by the addition of specimens of an
age group not represented in a series of the most frequent element.
For example, if the most abundant element of a given species in a
level was the mandibular ramus and there were three right adult rami and
one left juvenile ramus, the minimum number of individuals would be four.
Data on the modern fauna are derived largely from the literature
and from my notes on Trinidad mammals. In the course of my own field
work in Trinidad, Recent mammals were collected and their ecology
studied, with emphasis on those forms which are of importance in mid-
dens. Smaller species were collected in traps. Since the large game
animals are very scarce, I accompanied professional hunters, and acquired
the unused parts of their catches I also obtained many bones, par-ti-
cularly skulls and feet of vild mammals, in modern kitchen middens
around the nuanterb nou~es. ParLIther data on Rece:nt cmamma~ls were
obtained through tne analyrsis Of Ov1 p~lletB COllected ty trre late
F. W. Urich of St. Aug~ustlne and lent. to me byl Dr. H1. G,. Kugler aid
Dr. S. Schaub; fromr the Basel INatural Histojry Museum.
where adequate series were a-vailable, comrparisons of skceletal
moeauremnfts of midden and Becent rrammals of the sameP species were
undertaken with the aim of detecting differences in size that might
perhaps be correlated wit~n ecological or other factors.
POST-PLEISTOCENE HISTORY OF THE GEOLOGIC,
CLPIATIC, AND VEGETATIONAL FEATURES OF TRINIDAD
For the understanding of the history of the mammalian fauna
of Trinidad, a knowledge of the post-Pleistocene geological events,
and the climatic and vegetational changes attendant upon them, is
of primary importance. A brief sketch of these changes will, per-
haps, aid in placing the history of the fauna of the island in its
Although Trinidad is commonly thought of as one of the Antilles,
and is politically a member of the West Indian Federation, it is geo-
logically and biologically closely associated with the South American
mainland. The island is composed of sediments probably derived prin-
cipally from the Guiana shield which extends roughly as far south as
the present frontier of Brazil and includes Trinidad, and possibly
Tobago, at the northern limits of its influence. According to Dr.
Kugler (personal communication) several Pleistocene marine terraces
are traceable, and he has recognized at least seven of these, rang-
ing in elevation from five to 330 feet, which can be correlated
closely with similar marine terraces in Venezuela. After the last
glaciation, there was one continuous land mass encompassing what is
now Trinidad, the Gulf of Paria, the Serpent's Mouth, and Venezuelar.
Thereafter a general trend of rising sea level is observable. Kolde-
vijn (1958) states that the shoreline at an estimated 17,820 years
ago was at the present 24-fathom line (Figure 1). By about 8,000
years ago (Figiue ;), the ;thoreline had risen toc thle present 12-f'athom
mark (Nota, 19?5d). FRoughly: 700 years ago, t~he drainage o~f the Manamo
Branch of the ijrine~co River intG the OulP Of Parih 10creased in VOlume~
thereby further enlarging thle Gulfr, and completing the s~eparatioo of'
Trinidad as an island from the mainlania (KoldewiJn, 1958). At thle
present time, the island is rubsldingI, as is evidenced by the cGasth]
erosion indicated by the landalips onto the beaches. May instances
can be seen of roads that have been completely undermined by wave ac-
tion and of coconut palms lying uprooted on the beach. The Chagornaray
site on the south :oast is being waEshed into the sea.
Prinidad may be divided into five physic~graphice regions (beard,
1946). The first of these comprises the mountatus of the Nocrth Ra~nge
(Figure 3). llthough there are three ranges in TIrinLdad only the
North Range, which is an extension of the coastal cordlilere of
Venezuela, may be considered to be actually mountainous, rrlth eleva-
tions over 3,000 feet. The Central and Sourth ranges constitulte the
second region. The elevation of these bills scarcely exceeds 1,000i
feet. Just south of the NYorh Range lies the third region, one of
dissected alluvial terraces, formed of m~aterial from the mountains.
The fourth and most extensivre region is the dissected peneplain.
This division includes all of thre rest of the land except for a fews
coastal swamps which comprise the fifth and final region.
As a result of its insular nature and varied physiographic
features, Trinidad has a somewhat varied climate. The most notable
Figure 1: The shoreline of Trinidad about 17, 820 years ago.
The solid line is the present 24-fathom mark which
corresponds to the former shoreline, and the dotted
line denotes the present geographic features of the
Figure 2: The shoreline of Trinidad about 8,000 years ago.
The solid line is the present 12.fathom markr
which corresponds to the former shoreline, and the
dotted line denotes the present g;eog;raphic features
of the area.
Figure 3: The physiographio regions of Trinidad.
climatic zone is the montane region of the North Range, which is
characterized by heavy rainfall of at least 200 inches per year and
an almost continual cloud cover. The remainder of the island experi-
ences a seasonal climate, with two dry seasons from January through
April and September through October, and two rainy seasons, as the
result of shifts in the tradevinds from northeast to southeast. ThZe
windward coastal strip is affected by the vind and salt spray, and is
therefore somewhat drier than the rest of the island. Rainfall data
for selected stations, provided by the West Indies Meteorological
Service, are presented in Table 1.
Evidence indicates that temperatures in the area of Trinidad
underwent an increase, beginning about 16,500 years ago, reaching a
maximum at 6,000 years ago, and thereafter declining slightly
(Exiliani, 1955). The enlargement of the Gulf of Paria may have had
a moistening influence on T~rinidad. Present day temperatures vary
in the different parts of the island, but on the whole are generally
equable. The temperature ranges at Port-of-Spain, representative of
the seasonal climate, vary from the average monthly maximum of 86.4o
F. in January to 89.80 F. in May, and from the average monthly mini-
mum of 67.60 F. in February to 71.80 E in September. Daily tempera-
ture fluctuation is almost constant, ranging from a maximum of 18.90
F. in February to a minimum of 160 F. in September. Humidity always
exceeds 50 percent and often approaches 100 percent at night (Beard,
.--1~~~ u' O ** Pa l
'.U .3 5 8
H ad rT i
S20 O9 0 CO U .
P u 1 ,n C F
ah us 01 st.
7L C ;rld d =
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n d -,-e a
rb oj m8 e 3 =
Striking changes in the vegetation of Trinidad have accompanied
the geologic and climatic events that have influenced the island.
The following discussion of the vegetation is based upon Beard's
monograph (1946) The Natural Vegetation of Trinidad. At the time
when there was a continuous land mass including Trinidad and the
mainland, there was also an extension of the vegetation types of
Venezuela into Trinidad (Figure 4). The North Range was forested,
as were some of the higher parts of the Central and South ranges,
while the remainder of what is now Trinidad and the Gulf of Paria
vere occupied by savannah. The savannah was dissected by streams
bordered by gallery forest. The savannah formation which is now wide-
spread in Venezuela as the "llanos," reaches to within 50 miles of
the Gulf of Paria and persists as relics in a few small areas on
Trinidad itself. The distribution of savannahs on Trinidad is con-
trolled by edaphic factors. They occur in flat areas where the soil
is poorly drabed and the subsoil is impermeable. The soils became
rejuvenated, accompanying geological changes in the area, and this
resulted in the spread of forests from the streams and hill, eventu-
ally to cover nearly the entire island.
Particular climatic and edaphic factors influenced the develop-
ment of several vegetational types. Beard (1946) recognizes six major
plant formations (Figure 5); seasonal forest; dry evergreen forest;
montane forest; intermediate forest; swamp; and marsh. Four of these
are further divisible into associations. The seasonal formation
inclu.des evergreen, semi-everrreen, and deciduour forest depending
upon the aVailability of moisiture.
Th~e everqreen Peaso-nal forest requlreE ;C0 inenrte or mo_re of
avahiitlabl moliture, and3 is CharaTcterized bry a dioco~ntinuousl e-mergent
jtratum mor~e thanr 103j feet in neight,~ and .. an almost contiou-
ousI canopyi ilaer at Li0-':i feet, a~nd almostr co~ntinuous~ lower storry at
10-3,0 feet" (Berdr~, 1966~, p. 38). LI~nua and epiphlyter are scmmon.
C'onspicuousl IpeciFes ofr trzee are 3ponding csrmbin (hogplllum) Carype
eulanen~is (CraFPpo), Eschue~llrer sutblandalora ((quaIteCare), Panta-
clethra macrolota (Dol- ma~latra or fineleaf), Maximilinan elezansj
(cojcorite palm), and Mol~rre exelsr (moira). The 1largest tree s re
bt~tressed,~ andj mny are imp~ortant :oaruces of food for uild aninals.
Phe EeCi-Pev~Ergen Eeasoal4 forest exii-ts '-here 50-70l inches
of available moisture occurs. It is an open fo~rest characterized by
...a diEC'ODt~ilIDUlE ewrTgetjL le8er 80 tOJ-cq fret,,j *>D 80 81OSt
continuous~ later qt go.LO feet (B~eard, i1?66, p.. 3!'). Lianas are
very abundajnt but epiphytes 4rE not. Buttrefced tree; are rare.
Commiiron pFecies are Biravastal integerria h (jiggler-ueed), hnd dura~ crepi-
Ene third form1I of tEasonal1 foreEt l thie jeciddlcus,. OjccOT3 in
areas of !0-50 inches of' available moistu*re. It 1: Chharaterized
byr naving~ a co~ntinlUGUE~ canopy at 10r-!0 feet, aind a f'ew emergent trees
up to t60 feet in heig Lt.
The, dlry evergreen o~r littoral woodland formation oc~cure alongr
the coastal Itrip which if EXpoitd to continual wind Off thr oceRD.
Figure 4: The Pleistocene vegetation of Venezuela and Trinidad.
Figure 5: The pre-Colutibian vegetation of Trinidad.
Two associations of this formation exist in Trinidad: the Coccoloba
uvifera-Hippomane mancinella (sea grape-manchineel), and Roystonea
aleracea-Manilkrara bidentata (palmiste-balata).
Beard divides the montane formation into three association:
lower montane rain forest, montane rain forest, and elf in woodland.
These replace one another at successively higher altitudes. The
height of the canopy decreases with increasing altitude, from 70-100
feet in the lower montane rain forest, to 60 feet in the montane
forest, to 15-25 feet in the elf in woodland.
The intermediate or seasonal montane formation is intermediate
in characteristics between the seasonal and the montane formations.
The swvamp formation is composed of four types: swamp forest,
palm swamp, herbaceous swamp, and masngrove woodland. The greatest
area of swamp forest is the Oropouche swamp. In it the principal
tree species is Pterocarp~us officinalis (swamp blackwood), which form
a closed canopy at 60 feet and is conspicuously buttressed. The
species of palm occurring in the palm swamps near the sea is Ro-
stonea oleracea. The marsh formation consists of marsh forest, palm
marsh, and savannah. The swamp and the marsh formations develop
under the influence of particular edaphic factors. The marsh forma-
tion is characterized by only seasonal inundation and the swamp
formation by permanent inundation. Savannahs of the marsh forma-
tion are characterized by a dominant short bunch-grass, and a scat-
tering of gnarled shrubs attaining a height of up to 12 feet. Several
species of grasses occur, and the chief shrubs are Byrsonima crassifolia
(.avann6 -erratta) and CI1ratella emericano (rougihlear) or c(hrlso-
balanius pEllocarpus (f'at polri). Inere fire-resistant -.hnabe have
in onez areas replaced thec formerr Myrcl-ria-ouae ascsoation Meni
:till GCCufrE vwidly in the Venezuelan 11ano;.
The natulral condition of the- vegetation a ir.etchled above
has been markedlyr modified in recent times by the agricu~ltural
activities of osn. About half of the Ilin~d is still forested. Tne
land that 1. ulnder cult~ivation use forme-rly: prrinciplly ealconal
forest. Cocoa and COlffee are grson m'iinly~ in the ;lorh aind Central
Eng~er, unereas sugar cane la grown in the flatter areti between
these ranges. Rice is culltivatedl to the 'camp areas and co~conute
along the coast. Farticularlyr the east cojast. There has been some
reforestation Lcith teak~ in thie southern Ystershied.
THIE PRE-HUMAN MAMMALIAN FAUNA OF TRINIDAD
Phe samples of the mammalian fauna from the three different
time periods to be treated will be considered separately. The old-
est period, represented by fossils, may be designated the pre-human.
The Recent fauna, representing two periods, is composed of Indian
midden bones and of specimens from a period that may be termed the
historic. The time range for the pre-human period includes the
Pleistocene epoch and probably the early post-Pleistocene as well.
A carbon-14 date of greater than 34,000 years has been determined
from wood associated with Megatherium remains at a depth of 12 feet
at Trinidad's most important fossil locality at Forest Reserve, the
so-called Glyptodon site. During this time Trinidad was still broad-
ly connected with the South American mainland, and savannah dissected
by streams edged with gallery forest covered the area.
The pre-human fauna of Trinidad known at present is small,
reflecting the limited amount of collecting carried out in thi-; field.
The fauna is represented by three orders: Edentata, Rodential, and
Proboscidea. The edentates are represented by the families NMega-
theriidae, Myblodontidae, and Glyptodontidae, the rodents by the
Cricetidae, and the proboseidians by the Gomphotheriidae. All of
these families, excepting the Cricetidae, are now extinct.
Fossil Mammals of Trinidad
This family is represented by Megatherium americanum Blumb.,
whichh he beenj fcud at 3Pveral localities.C AO 5Stregalus1 was
found at Kasramat mud voicsno and identified tiy Schbsb (1035E). Parts
of r js:eletonn wre foulnd at Lo~s BaJe-e~. The site at For-'str Reierve
alrco yicloed IMesrthrrium remalcin. Pa-rt of' a toctch found at the rlmi
the Jlypted~on~ ;i?-eleton wra e:;cavter d (195)~;, ird along with the other
masterials, Yas ;ent ro the Americran Mu emw of Natural History. In
1959' I f~ound manyG diErmal becnes, a complete foulrtrr tersal, a complete
tibia and fibrlls, part ofi s mandibullar rtimus, a rib, and tnree
thoracic vertebrae. Tn~is fosail atte is by far the muost. pro~du~crtve
on Trinlidd. It ii a ri'.er bed cutr to~ a depth of 15. feeT itotc
Mi~cene: IAnds and claysj, and~ filled with~ oil impregnatedj Ilt, rand,
clb; 3ttOe pebbles, plant remains rangilng fromr large logr tc. tuiE-
and stcmj, and vertebrate and insect fosaill. TInla site is, knownr
1iccelly as the 'Glyptedon eite."
& alngle too~th Or' IMV10duoD Ep. .E C-Xcava'tCd along Uith othrbc
materials at the "Jly~rtodon jite" and has been identified by me. It
ii notr deposited in the collection of the America-n MuseusLL oI if NtraL
An elmot. coeplet~e ilyptedojn ateletcon, from which t~he 'Glyp~tG-
do~n iite' takes itj namE, we5 found whi~nle the oil vell site was be-
ing prepared in 195;. Almostr the entlre si-.eletn wa found In place,
and is rlu, un~prepared, at rhF American MI:eum o~ Naterurl History.
I found abundant dermal bone but no other skeletal parts while working
at this site.
At the "Glyptodon site" I found a layer with a concentration
of small vertebrate remains two feet above the layers contain-ing the
large bones and logs. A single mammal was represented which is refer-
able to the genus Zygodontomys. Associated with the Zygodontomnys are
unidentified remains of a few birds, a turtle, a frog, and many in-
sects. Another site containing small vertebrate fossils is at Apex
near the "Glyptodon site" and geologically similar to it was found
by Dr. Kugler. Zygodontomys sp., the only mammal represented, is
associated with bird and insect remains (Blair, 1927).
Named species of the genus Zygodontomys are distributed in
northern South America and southern Central America. Forms of _Zygo-
dontomys have been described from south of the Amazon River, from
northeastern Brazil and the Nbtto Grosso, but these are included cor-
rectly in the genus Akodon (Tate, 1939). Tate (1939) divides the
Zygodontomys into two primary groups of small and large size. In-
cluded in the group of large-sized species is ZyEgodontomys brevi.
cauda, which occurs abundantly on Trinidad, and other species dis-
tributed in Panama and northern Colombia. The group of small-sized
species include ones distributed in Central America, Colombia, Vene-
zuela, and the Guianas. Tate (1939) also reports that among the
specimens referable to Zygodontomlys microtinus stellae from the Es-
merelda savannahs at the foot of >k. Duida there are some individuals
rith a normal tipper molar tooth row of Ir.T' a. and there are others
with the shorter knwnm tooth row of 5.7 to j.8 mlm. for tnLa genus.
G~yldecrstolps (193~) notes that Sygodo~ntomlr thomast distributed in the
Cumansrr district of Vlenezuela 1: allied to ygodontomys brevricude but
ie much smaller. The focsil LyodontoySE from Trinidad i; also siclar
to iZivgodonrtemprr s brevicanda6 but a great deal smaller.
Since the Cossil material is largely Trcrmentary only a few
mbEauremen~ts could be taken. The mojst 112eful ones were the alveolar
length of the uIpper and lower cheek tooth rovp A comparable ;et of
measurement; VEE ta.4en On ZYRj00Odtmys tlrevicaudaB from oVl pellets
found by F. 'd. Urich on PInidad, Zygodontomya~ micro~tinue stellae from
Bameralda Salvannah, Venezuela, and "vgodotomei: tnomast~ from Cristobal
Colon, Venezulela. These were taken to the nrearest one tenth of a mm.
and appear in rtale 2.
The fos-,il Spodorrtomyr diff'ere from th~e other species of "ygodon-
tc~ougg available for cojmparison in eeveral respects. T~he most, outstan~d-
105 characteristic of the fossil form Is Its small size (Teble i). Ad-
ditional cararcteristics that distinguich one fossil are exceptionally
long anterior p~alatal foraminra and deeply concave eygomatic plate. In
the younger individuals (specimens with unborn teeth) the foramina extend
poeterior to the level of the posterior lingual root of the first molar.
In the oldest individualP of thre f~osel (vith heavily vorn teetb1 and
in young individuals of other rFpeciet the foramina extend posteriorad
only to the anterior root of the first molar. Diagacstic dental charac-
tertatic include a deep anterior median fold in MI, a broad U-shaped
MEASURIIEHTS (IN MILIMGTERS) OF THE ALVEOLAR CKEEK[ TOOTH ROW
OF VARIOUS SPECIES OF ZYGODONTOMYS
Z. microtinus stellae
I fr I I
* 5 a ; Co
i, i; ru a
I I. ~ I
f- *1 CC
first primary fold, and presence of a mesostyle, anterior labial style,
anterior' lingual style, or enterostyle in the M1 and M2. The anterior
median fold is exceedingly well developed and is present in all speci-
mens except those in which it is obliterated by excessive wear. This
fold is otherwise present in only a few specimens of Zvgodontomys brevi-
cauda and in these it is merely a vestige. The first primary fold is
broad in the fossil specimens and narrow in the other species. In the
MI the mesostyle is present in all of the specimens and the other three
styles occur in between 64 percent and 73 percent of the 16 specimens.
In the M2! the enterostyle is present in 78 percent of the nine specimens
and the mesostyle in 67 percent. No styles are found in any of the other
species. In the lower cheek tooth series the mesostylid is present on
the M2 in 94 percent of the 17 specimens. The articular process of the
mandible is also distinctive, being short and broad, whereas in the
other species it is long and narrow.
According to Tate (1939), "The South American Zygodontomys ap-
pear to be a g;rassland-inhabiting group of mice. I have little doubt
they extend southward from Sucre across the 11anos to the Orinoco and
continue into Guiana wherever savannas exist." It appears, therefore,
that when the 11anos extended into Trinidad there was opportunity for
the spread of this genus into Trinidad from South America.
Cuvieronius hyodon (Fischer) is represented by two teeth found
at Los Bajos and identified by Schaub (1935), part of a tooth found
originally at the "Glyptodon site," and another fragment of a tooth
found by me at the same site.
~oogeographic Signifleance ofP Pre-human Fauna
Ene known fossil faunae of Trinidad is sperce in numbers of
species. Further collecting will probably increase this number con-
siderably. The Edentates are of South American origin but the three
genera, Megatnerfum, Mylodon, and Gllytodon, found in Trinidad were
distributed througout much of both North and South America in the
Pleistocene. ~Apptherium ranged frcn South America to southeastern
United States in the Pleistocene. Bylodon ranged from South America
to southwestern United States and was contemporaneours with man. Glyp-
todon was apparently very numerous throughout South America and
southern United States in the Pleistocene. Of the Proboscidea, the
genus Cavieronius_ probably originated in southwestern United States
(Osborn, 1936) and was distributed widely in South America and in
southern North America in the Pleisto~cene. The rodent Zygodcentoips
evolved in South America and exists now in South America north of the
A~mazo~n Piver and in Central America as far north as Costa Rica. Certain
forms south of the Amazon River which havre been referred to the genus
are probably referable instead to the genus Akodon (Ta~te, 19?3). All
known representatives of the fossil fauna of~ Trinidad are herbivores
and are characteristic of greasland haBitFtats, which vere widespread in
Trinidad during the Pleistocene.
MAMMWALIAN FAUNA AS REPRESENTED IN INDIANI MIDIDENS,
The present knowledge of the pre-Columbian mammalian fauna of
Trinidad is based on skeletal material excavated from nine Indian
sites. These sites represent all the cultural levels known on Trini-
dad. The sites, like all those known from the island, are located near
the coast, indicating an orientation of the people toward the sea.
The pre-ceramic sites are, of course, the oldest, dating here from
2,750 (+ 130) years ago. These are followed by sites exhibiting
four distinct ceramic styles (Bullbrook, 1953) that span the time
from the pre-ceramic period to contact with the Spanish after 1532.
The first of these periods is characterized by the Cedros style,
the second by the Palo Seco style, the third by the Erin style, and
the last by the Bontour style. Pottery of the last period is some-
times associated with Spanish ceramic work (Table 4).
Description of Sites
1. St. John
The St. John site is located on a high bluff south of the
Godineau River above the Oropouche Swamp, and at the end of the St.
John Road. This location takes advantage of the evergreen seasonal
forest on the hill and the mangr~ove swamp habitats. This site was ex-
cavated in 1953 by Drs. Rouse and Goggin (Rouse, 1953). The soils
are light sands that are freely drained. The shell midden at this
site is extensive, reaching four feet in depth and 125 feet in dia-
meter. Material of two successive occupations has been obtained
CORRiFE1)1lONl ;J PERIiOD OF ~:cCUPAT~ICn OI ITR~LD;lDA AND~ CEhrAIC JTIYLEi
afterr CM-entbt and PouC.)
Vcnezuelan Ekriods of Tr~inidad Ceramic
Occupnt ion ttyle;
? St. Jee~ph~
here. The first of these is pre-ceramic and is comparable to the
Ortoire site, for which there is a carbon-14e date of about 2,750
years. The second period of occupation is characterized by pottery
of the Bontour style, with :rome European vare. This excavation con-
sis~ted of 13 level units, four of which were pre-ceramic, five proto-
historic, and four a mechanical transition between the two. Each of
these three groups of levels has approximately the same proportion
of each mammal species in them, but the pre-ceramic group contains
more bones (Table 5). This is probably the result of differences in
the economy of the two cultures, the pre-ceramic population depend-
ing exclusively on hunting and fishing, while agriculture also con-
tributed to the food economy of the protohistoric population. An-
other factor that might explain the difference in the relative quan-
tities of bones and artifacts could be that in the ceramic sites a
larger proportion of the bulkier artifacts would correspondingly re-
duce the number of bones found in a given area and level.
List of Species Identified
Didelphis marsupialis, opossum: 10 individuals.
Casypus novemcinctus, nine-banded armadillo: 27 individuals. As in
all other sites there is a far higher proportion of bone fragments
to minimum number of individuals for the armadillo than for any
any other mammal because of the numerous bones in the shell. On
the average for every 4 to 5 individuals, there were over 75
armadillo bones identified, but fever than 50 bones identified
for the same number of any of the other mammals.
Frecerelmic M:echanical Prot~ohistoric
D~i d lFh i i 3 3 4
Ea iypus 15 6 6
AUlouatte S ?
Coendu 1 r' 1
Agout i 11 7 .
De cp roc ts B 3 d.
E: ni m:3 re2 1
ProachimyE 4 2 1
Pzcart 2 18 12
iELATIVE ArBUNDAN41CE iF ~Mr'dBAD AT D~IFFE~REWI'
ITEDJ~TRE 3T. JiOUN~ 3ITE
Alountta seniculus, red howler monkey: 10 individuals.
Rhipidomys couesi: 1 individual.
Coendu prehensilis, porcupine: 4 individuals
Agouti paca, paea: 26 individuals in all; 8 in the proto-historic
group of levels, and 11 in the pre-ceramic group. The remaining
7 were found in the levels of mechanical transition.
Dassyprocta agut., agouti: 13 individuals in all; 6in the proto-
historic group of levels, and 4 in the pre-ceramic group; 3 in
the transitional group of levels. The paea and agouti vere repre-
sented numerously enough in all sites to allow useful comparison.
Echinys armatus, spiny rat: 4 individuals.
Proechimys guyanneasis, spiny rat: 12 individuals.
Procyon cancrivorus, crab-eating raccoon: 3 individuals.
Lutra enudris, otter: 1 individual.
Felis pardalis, ocelot: 1 individual.
Pecari tajacu, collared peccary: 57 individuals in all; 12 in the
protohistoric group of levels, and 27 in the pre-ceramic group;
18 in the transitional group of levels.
Mazama americana, brocket: 24 individuals in all; 7 in the proto-
historic group of levels, and 10 in the pre-ceramic group; 7 in
the transitional group of levels. As with the paca and agouti,
the peccary and brocket were sufficiently well represented at
each site to allow comparison.
Other vertebrates: As at all other sites there were very few birds
represented, Cairina moshata (muskovy duck) being identified.
Ther werc a Tfew lend t~utles, butt no sea tuirtle remains at thiz
ite. Fion remains, particu~larly cat fi:n, were abnt~iidat bere a;
at all other -ites.
The Cedrosj site it ritua~ted on thre United -`rtate Arm's
Green H~ill Reservation in the extnrree coutrrwest corner of the 12-
land and is located on a rliynt elevatico uitniar one fo~urth mile of'
the sea. The area is nowu a coconut plantaticon and 1: inhabited; it
was, however, formerly Ielmu Jrsp surLrounded Dy evergreen saaonal
forest. The sandy scLL in this srea 1: classified as~ having Impeded
drainage, since it is close to tea level and the uater table. Cedrea;
is a shall mound yielding purely Cedrose style pottery. The materials
studied were from an ex.avaittoo made by fr. Rou e snd Mr. Eulllbrootr
(RomIe, 1953), consisting of 25 level unite, and from tio random
surface collections madle of ir. Eaglcr sad byr me. The eco~nomy of'
this culture, as is; trule for ill the ceramic cullturres known c~n Trint-
dad, was probably as~ed on bunting, fizbing, gathering, and agricul1-
List of Species Ideotified
Didelphis maraupialls, opossun: 5 individuals.
Egrygu+ ocY~vemeclletus nine-banded armadillo: 15 individuals.
Taimanduie Leagicelaudatanot eater: ;' individuals.
Alouatts :-eniciLlus, red howler monk~ey: 1 in~divdidual.
agogti Esac, paco: 16 individual;.
Emlrprous EEE11 goultt: Ilb individuls.lr
Proechimys guyannensis, spiny rat: 1 individual.
Canis ef. familiaris, dog: 3 individuals. These were represented
only in the surface collection, and were undoubtedly added in
Procyon cancrivorus, crab-eating raccoon: 2 individuals.
Herlpestes auropunctatus, mongoose: 4 individuals. This species,
like the dog, was found only in the surface collection and was
deposited in recent times.
Pecari tajacu, collared peccary: 14 individuals.
Mazama americana, brocket: 23 individuals.
Bovid: 1 individual. This was found only in surface collections
and was deposited in recent times.
Trichechus manatus, manatee: 2 individuals.
3. Palo Seco
The Palo Seco midden is on the south coast of Trinidad. It
is located on a ridge at an elevation of 90 feet and is surrounded
by hills. The shoreline is approximately 200 yards distance. The
Palo Seco site is now at the center of the Trinidad Petroleum De-
velopment camp, but it was formerly in semi-evergreen forest habitat.
The soils of this area are predominantly clays and silts with impeded
drainage. The excavation, made by Dr. Rouse (Rouse, 1953) at this
site consisted of 57 level units, and yielded two pottery types,
Cedros and Palo Seco.
Lit of Ipecies Identified
E~ddlphis marraupilzl, opossu~m: 1 individuals .
Egils. gy no~vemeinctus, nine-baneobd annadillo: 18 individuals.
Tramandui longicaudata, 6nt eater: 2 Individuals.
AloilnttB SCniC~ulus red howler monr.ef: : individuals.
Ccendu~ preheInailii, porcupine: 6 individu~als.
Agouti IaCa, paBce: b1 individu6lr-.
Casypracta. agyti, agoutli: 67 individuals..
Proechiwy: @.ayannensds, spiny rat: 1 individuals.
FE1Felis agjgliS, ocelo0t.: 1 dvi l
F.cari tez'gy:3 collared1 pee5ar: 20; Irindividulj.
blazamas hmericans, brocvtet: 76 indivlduals..
BGi:1 iodividuatl. Thlz *ras found at the -urf'ace, pre--umao9ly
from recent oc~cupation.n
Tsyling ip., tapir: 1iiiul.A rieht lowe-r recond pre-molar
was foulnd. Th~i s nimal ~cculrj on thef mainland of Soulta Aeric~a.
TRichechu; manatus~, manatee: 1 individual.
Cetacean : 1 individual.
ither ver-tebrates: ia tlUt~le remained er~c abundant aT. thls site~
and at the other coath coast ;Ite. mere were few or no reciains o
Che Brn midden Is Ju1 t rest of Palio reco in the center of the
townr of` Ern lnde~r thet present oilice ItitiCn.. It is on a hlill that
vaE CONered wlth Semui-eVergreDc rOretS, R't0ut One-half mile fromn tne
beach and the Erin River. The soils here, as at Cedros and Palo Seco,
are poorly drained clays and silts. TIhe site yielded Palo Seco and
Erin style pottery. The materials that were examined came from an
excavation made by Dr. Rouse (Rouse, 1953) that consisted of 69
level units, and one excavation made by Mr. Bullbrook, in which, how-
ever, the specimens from different levels were not kept separate.
List of Species Identified
Didelphis marsupialis, opossum: 18 individuals.
Caluromyrs philander, woolly opossum: 2 individuals.
Isusnovemcinctusnebaddamdl: 10 individuals.
Tamandua longicaudata, ant eater: 2 individuals.
Alouatta seniculus, red howler monkey: 3 individuals.
Sciurus granatensis, squirrel: 1 individual.
Nectomys s~quanipes: 1 individual.
Coendu prehensilis, porcupine: 4 individuals.
Agouti paca, paca: 14 individuals.
Dasyprocta agati, agouti: 59 individuals.
Ech ,,sarmtu, spiny rat: 2 individuals.
Proechimys guyannensis, spiny rat:2! individuals.
Canis cf. familiaris, dog: 1 individual. This came from a surface
Procyon canlctivorus, crab-eating raccoon: 1 individual.
Lutra enudris, otter: 1 individual.
Felis pardalis, ocelot: 2 individuals.
Hrpe-- es aulropunctatus,, ;ongoose: 1idvda.Ti aefo
Mlr. Ba~lle-rook'r :collection and p-res;umably wa~s collectedr fromI the
Pecari teggygy, collared peccary: 11 individuals.
Mazama americana, brockret: 61 individuals.
Trichechus manatus, manatee: 1 individual.
This site is just back of the Atlantic coast shore in a cocc~-
nut grove belonging to the St. Bernard Estate, a short distance di-
rectly south of Cape Mayaro. The ceramic style at this site is pre-
dominantly Palo Seco. Thne bones available for study consisted of
two small surface collections, one made by Dr. J. HIill of Port-of-
Spain and the other by me, both in 1959.
List of Species Identified
Da~sypus novemcinctus, nine-canded armadillo: 2 lndividuals.
Aaouti paca, paca: 1 individual.
LiCj7sypract aglt i, agoutt. 1 individual.
Procabinvrs ygyaonenhns, spiny: rat: 1 individual.
ecarl rta.jacyl, cojllaredl pccary: 3idvdas
Mazama smertcana, brocket: 2 individuals.
Ihe- Qul~ira aidden is located in semi-evergreen forest east
of the PFIlo ;eco site, near the Desch and the Palmiste River. The
soil ij similar to that of Win and Polo Seco. The excavations ruade
by Dr. Rouse (Rouse, 1953), consisting; of 58 level units, produced
pottery of Palo Seco, Eri~n, and Bontour styles.
List of Species Identified
Didelphis marsupialis, opossum: 4 individuals.
Igasyus novemcinctus, nine-banded armadillo: 10 individuals.
Tamandua longicaudata, ant eater: 2 individuals.
Alouatta seniculus, red howler monkey: 2 individuals.
Agouti pace, paca: 16 individuals.
Dasyprocta aguti, agouti: 26 individuals.
Lutra enudris, otter: 1 individual.
Felis pardalis, ocelot: 2 individuals.
Pecari ta geu, collared peccary: 23 individuals.
Tayassu pecari, white-lipped peccary: 5 individuals. This was the
only site where remains of this species were found.
Mazama americana, brocket: 77 individuals.
This site is half way between Palo Seco and Quinam on
Chagonaray Point. It has quite the same ecological conditions as
Quinam, but is undergoing coastal erosion. The pottery found was
principally of Erin and Bontour style, but also included some of
the Barrancas style typical of the Lower Orinoco region (Rouse,
personal communication). The mamrmal material examined was from a
surface collection made by me.
List of Epeie: Ilentified
D3egguy noveac~crincu ninc-banded arcadillo: 1invial
GEcul pae$, paCa: 1 Individual.
Desyprooctb agartt, ago~uti: 1 Ladividual.
ProePchings tuj-ouneoie, iFlay rat: 1 individual.
Pecari~ tagcu, .:ollar`Ed Peccaryi: 3 indirvidualf.
Mazama americana, brock~et: 2 individudls.
Masyc ii a Spanlish missior. rite locate-d inr the Miontjerrat
Hill orf the Central Range, Seven mils fromn the GuL~Clf of Pan. A
Catholic church haj been built over the site. Formerly this aree
waj covered byr evergreen Feasonal forest. The acoil is light and
freely drainea. On the btrsis of the bcentour style pottery andl the
Europeanr were foundl, this site has been dated (G~oggin, pers~nEJ l comm~u-
oication) from the late seventeenth o~r earlyr eigh~teenth ienturies.
The material examined cjne fromn an excsavaion ofP 8 levecl unitE made
by LT. Goggin (Rcouie, 19153) and frojm myr surface coillection.
Lirt. of Speciesj Ident~ified
I ~pus nov~encinctur:, nine-bandei armadillo: lb indi iduls.S
Tamaicndua longlc~audata, ant eater: 3 individuals.
AlUcatta seniculus, red houler monkey: 6 individuals.
Ccenda prenlnsliE3, porcupine: individuals.
Agouti pgya, paca: idvua.
Enerpracta aggil, agoutl: 5 individulalr.
Canis cf. familiaris, dog: 1 individual.
Pecari tajacu, collared peccary: 17 individuals.
Mazama americana, brocket: 8 individuals.
9. St. Joseph
St. Joseph was the first Spanish capital of Trinidad, and
the site is located in the town of St. Joseph on a branch of the
Caroni River on the south side of the North Range. This area was
semi-evergreen forest. The excavation made by Dr. Rouse and Dr. and
Mrs. Goggin (Rouse, 1953), consisting of 13 level units, yielded Bon-
tour style pottery and European vare.
List of Species Identified
Coendu prehensilis, porcupine: 1 individual.
Agouti paca, paca: 1 individual.
Dasypracta aguti, agouti: 3 individuals.
Mazama americana, brocket: 2 individuals.
Bovid: 2 individuals.
Cetacean: 1 individual.
.r '1 'i
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ir ~I le
rd .4 .5 "g "a ,-< 4 0 fi 0
32 .-- r ."a
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The mammals that have been identified from Indian middens com-
prise a surprisingly complete list of the larger animals found in Trini-
dad today. One group of animals that is, however, entirely missing is
the bats, although there are a great number of bats (Goodwin and
Greenhall, 1961) now living on Trinidad. Another group, the cricetid
rodents, is represented in the middens only by the two largest genera,
Rhipidomys and Nectomys. Of the four species of marsupials now occurring
on Trinidad, two species of the genus Marmosa were not found in the mid-
dens. Presumably the bulk of the animals represented were used by the
Indians for food. This automatically excludes from the middens a cer-
tain portion of the natural fauna of the area, either because the Indi-
ans would make no effort to catch animals they considered useless, or
because they might not have the technological ability to catch other-
wise edible forms. Thus, for example, although there seems no reason
to suppose that the bat population on Trinidad was markedly inferior to
that at the present time, it is not wholly surprising that no bat re-
mains are to be found in the middens.
The smallest animal represented in the middens is Rh~ipidomys.
With three exceptions, all larger animals now known from Trinidad occur
in the middens. One of those not represented is the small arboreal ant-
eater, Cyclopes didactylus. This animal, aside from being relatively
small, emits a very mournful wail, which may have caused it to be
avoided. The capuchin monkey, Cebus albifrons, was also not found.
This species has a large gap in its distribution on the mainland, and
the cloieat 4lly to the Trinidadian form occu~r s round th Maracaibo
Ba:1ln in ehstern~ Venezuelai and nowhere in thie arecs clo-Fr to Piknided.
HErahaevit:= (134?) hi; suggestpd two po_lible eplanations~ focr its apF-
eanrance on Prinidad: that Ctbut. altifrons 64ay at. an earlier time havf
nad a uider diatriaution trhan it dec. at present, or that it was
brought tor Trinidad by early Irnhabitant; a3 a pe~t. in regard to the
second of these aggePSt~ions, it Beem: hitghly dOubtful* that, had the
monkeyr beon kept ab~out the camrps, no remains~ of it *ould be found. The
absence of' Inli PpeCiSJ from the midden rmaterial does not, howuever,
Ieo nece.sarly tc. invalidate the first suggestion that it Las In-
dgernous to Prinidad. Posalbly the explanation lies in thle faict that
the aonkrey is of relatively smcall s~ize, and may~ have been unde~istirbe
as an artic-le of fi.ood. The third animal abilch might havIe been expected
tut uset not recorded i; the tayra, Eira bFarbare. Rovever, rremain of it
have~ been recordedi by Bullbroot: (19?53) from n15 excsavatrion l at Pale 3ec.
Becainrs of' tuoc other large Fanimals whion may exist on Prinidad--
the -osti-munmdi, Najuh, and tne Iwater opo~ssum, Cairaectea--vere also
absent from the middene. Probably Individuarls of these species find
their way over from thre mainland from time to time. bh*. L. Venekind
(personal communications) ba; informed me that thle coEsti-aundtr possibly
occulre in the North Range. There 1: a somevbat better record of the
vuter opossum (Greenhall, 1956,) based on a photogbraph 4nd the descrip-
t~ion of a strange opossum1 shot by a hunrter inCla Guya~uayare at thie sou~th-
eastern tip of Trinidad. Phe sabence of remains of these an~nimle In
the middene, taken together with thE~r present status on Prinided, would
seem to indicate that in the past as now they were either merely oc-
casional visitors to the island, or, if native, so rare as to be of
no consequence to the fauna.
Only one mammal not previously reported from Trinidad was repre-
sented in the midden samples. This is the tapir, Tapirus, of which the
remains found consisted of a single premolar. The tapir now exists on
the mainland, and it is quite conceivable that the tooth found at Palo
Seco was brought there by man. The white-lipped peccary, Tayassu pecari,
has been reported from Trinidad, but it is questionable whether it still
exists there or ever did in considerable numbers. Remains of it were
found at Quinam. Another mammal, the savannah deer, Odocoileus gymnotis,
which does not at present occur on the island, was identified from the
Erin midden by Dr. S. Schaub, according to a personal communication by
Dr. Khgler. I found no remains of this animal, however.
The surface levels at the Cedros, Palo Seco, and Erin sites con-
tained remains of introduced animals that were probably deposited at
the site subsequent to occupation by the Indians. Each of these sites
is in a populated area, making this supposition even more likely. At
Mayo and St. Joseph, as a result of colonial contact, these introduced
forms were incorporated in the middens. Bovid remains appear in the
Cedros, Palo Seco, and St. Joseph Middens. The mongoose, Herpestes a.
auropunctatus, that was introduced into Trinidad in 1870, occurred in
the Cedros and Erin sites. Remains of a small dog, Cania familiaris,
were found at Mayo and Cedros.
In Trinidad, comldering; all sites togetuer, -5 specie. of
moc~amal are rep~rreseted (Tatle o). Much can be learned- from those-
species whlich vere caugEht in Dsuffi~ient quasntitie s to marke them, on
the basis of the evildcnce of thie IireaInsJ found in the midden;, the
chief sources of aseatt to the Indian'; diet. In every ;ite the five
most abundant ;Fpecies wrere the 4rmadillc, Iij;;rus novemainrtuj; parea,
EEEE11 Eoc'a; agouLti, EssvprPocca agati; coillared PeC'ant,~ Pecari tu aglu;
and brocket, Hazama iameri-ana. Theise five forms make up~ more than 75,
percent, in number of individuals, of the aonimas ht each of' the s~ite;.
The incidence of thiese formrs at the six largest sites was anhllyzedl
statistically for rahndoness by a contingency table. Thre results
(x- = 115,.O, 4 degrees of' freirdom) shGH tbht the Ob~er~ed difr'ereDCOB
are highly sigJlificant.. There arer two possibilities to explain this
difference; either food preference by: the Indisns or the 10081 htimal
population. ;incp .here is no cu~ltural evidncerc to Irdicate a differ-
ence in food preference, thje co~nclusion that is drawn is that the three
ismaller one;, the Rrmladillo, paca and agouti, and the two larger ones,
tile penrys- and the brockett were probably equally sought. Therefore,
the relat~ive abun~dance of each at the different ;ites probably reveals
their aCtual iabundance in each area rather than selection by the Inodians.
At the three Iouth coast sites of Quinamr, Palo Leco, and Eria,
the percentage of armadillo among the mammale represented ulthiln each
site is one tjalf or one third as great as within the sites of M~ayo, 5t.
John, adL Cedre~s (see Table t6). .this distriburticao mny well be correlated
with one soil best suited to the burrowing activity of the armedillo.
At the three south coast sites, the soils are predominantly clays and
silts with relatively poor drainage. Thus soil conditions would not
be as favorable for burrowing as light soils with free drainage or ex-
cessive drainage found in the area of the Mayo and St. John site. The
sandy soil at the Cedros site, although classified as having impeded
drainage because it i-; close to sea level and the water table, would,
however, provide suitable substrate for burrows. The evidence suggests,
therefore, that the relative abundance of these animals is related to
the character of the soil as indicated by midden remains.
The relative abundance of agouti and paca at the various sites
would also appear to be correlated with the ecology around each site.
At the two sites in the denser evergreen seasonal forest--Mayo and St.
John--paca were found to be approximately twice as abundant as agouti,
whereas the reverse was found to be true at the three sites--Erin, Palo
Seco, and Quinamn--located in more open semi-evergreen forest. At Cedros
near evergreen forest, but actually in palm swamp, paea and agouti re-
mains are almost equally abundant. The paca is characteristically a
forest animal, feeding on fruits and eggs laid on the ground, although
it does venture into the edge of clearings as well. The agouti is usu-
ally found in more open forest.
A similar pattern is seen when the relative abundance of deer and
peccary remains are compared. At the Mayo and St. John sites, the re-
mains of peccary are more than twice as abundant as those of deer, where-
as at the Erin, Palo Seco, and Quinaml sites, deer are almost three times
as abundant as peccary. At the Cedros site also, more deer than peccary
remain wrere found. The Qulinam silte revealed a fev t~onr- referatle to
thrE vbite--lipped peccSIry, Tagaga! pecari, as "ell as5 remains of the cot-
laredJ peccary. Pa~ccary are often found in forreste area, and par~ti-
Cula~rly. along river bottomm, rwhere their feeding babits are similar to
throse of pigs (Seton, 1029q, quFting Auduton and Bachman). ideal habritat
of this kinLd is present at the Clt. John -ite. Phts habitat aas alsoi follnd
at bbyar vith the Ma~yo River providlyg the river bottom conidit~iou. Lear,
althoughn numrc~ous in denselyr foresed~ areac, prefer the open forestE
foulnd along the south cast.
Alith the Exception of the St. Joh site, obLch LP a 3pec~id caseI
no -tgnilficant treojs in the numlbcer of individuals of a given specie;
at different levels of excaivatiojn in the other sitesa could be discovered.
Mloreov r, vben the faUnas from the vaCioUI Indian middenas re compared
on the basis of the total f'auna rather than by:, levels, no striking dif-
fercnces are found in their compositioo. ;'nimals that are found only In
a few suites appear to be incidental, and are represented by only, a bone
or two. As nas been noted, differences in the faunse can beF observed
in the relative ab~undance of certain forma unio-n were locally n~umerous
and presumabl,. equally bought after by~ the Indianl.
HISTORIC MAMMAL FAUNA
Due of the first lists of Recent mammals of Trinidad was pub-
lished by De Ver-teuil (1884). It included 24 species of large, con-
spicuous land mammals. Three of these--Gulo, Viverra vittata
(= Grison vittata), and Cachicamus se~ptemcinctus (= Dasypus septem-
cinctus)--have since been found not to occur on the island. De Ver-
teuil's list was followed in 1892 by a preliminary list of 24 species
published by Oldfield Thomas. This number of species was enlarged by
the workr of J. A. Allen and F. M. Chapman which was based on material
collected in Trinidad by Chapman in 1893 and 1894. In their first pub-
lication (1893), these authors listed 37 land mammals, 13 of which had
not been previously reported. New species that were described included
Nectomys palmipes (Holochilus squamipes listed by Thomas is later
synonymized with this), Tylongs couesi, Oryzomys speciosus, Oryzomys
trinitatis, Oryzomy~s relutinus, Oryzomys brevicauda, Loncheres castaneus,
and Echings trinitatis. In the second paper (1897) they omitted
Mtyrmecophaga jubata, Choloepus didactylus, Cercoleptes caudivolvulus,
Loncheres castaneous (= Loncheres guianae = ciy armatus), and
Dicotyles labiatus (= Tayassu pecari). However, recent evidence has
been obtained to indicate that the last two forms do, in fact, occur on
Prinidad. Allen and Chapman also added the following species: Oryzomys
delicatus, Akodon frustrator, and Thylamys cardl. The most recent list
of Trinidad mammals is that of Veaey-Fitzgerald (1936), while the Chirop-
tera have been dealt with in a monograph by Goodwin and Greenhall (1961).
Cone f'orm included In the VPEsey-FitzgeraldJ list, AV.odon fra1 trato~r, evi-
dently~ ses described froa a juenle ;pecimen of' Typiccjontomy brevicaiuja,
ani had earlier been synonymized~ (Gyldenatolp~e, 1932). A seecnd speclie
given, Crlrj.omys breVICaudeI had' p~revIC'Slj beeno referred to thel genue-
2,'godontom;,s (Gyldeasqtolpe,, 103).
H list of the RePcent land rnammasi of Irlinided i.- presented below.
AlthoughP Dyr fr the largest .Pegrant of the mammei a~lr faua the Island?
iis mae up of batP. CI8 ageeles naving been reported, the Cntroptrer are
not inclu~ded in the followuing list, since they have been the subject of
a very~ recenrT monograph, andj 4re of conparatively little imiportance
archaeological. The liEL I9 e9960tla11] that of Vezcey-Fitzgerald ex-
cept for tne baddtiGO Of' two species, ana certain nomendaltural changeZ.~
Mosit of~ the nomnenclatural changes folilow the work of' Hershk~ovitz (1.:r7,
196~8, 1969s, 1~55, 1960) anil Cobrera (1P5;). En~e zsynogray for the
ipecies concerns only~ the references to the occurrence of' these jpecies
on Trinic~ad. The li.:t includesr observations that I made during the am-
rmer of' 1?5 .
L -t of Terrestrial Mamm~als
Tldelenlq manouoalls insularr~is lln, Bull, Am~er. Mus. Ijat. H-ist.,
rrldelphcB mereup~ielis Thcesas, Jiur. Trinidad Field Iati. Club, I,
Dide~~lphi mraeupelia Allen and~ C'hapron, Bulil. Am~er. Mus. Natir. H-ist.,
Didelphis karkinophaga Allen and Chapman, Bull. Amer. Mus. Nat.
Hist., IX, 1897.
The opessum or "Manicou" appears to be quite common, particular-
ly around plantations. One male was shot on 3 September 1959 on a cocoa
estate in the North Range, Arima Valley, Spring Hill Estate. Its
measurements were 744-399-55-En 48. Mango, insect and rodent remains
were contained in the stomach. One skeleton from Biche was obtained
from a bunter.
Caluromys philander trinitatis (Thomas)
Didelphis (Philander) philander Allen and Chapman, Bull. Amer.
Mus. Nat. Hist., V, 1893.
Didelphis (Philander) trinitatis Thomas, Ann. and Mag. Nat. H~ist.
(6) xIII, 1894.
"Manicou gros-yeux" is also quite common around plantations.
One male was shot on 3 September 1959 at Spring Hill Estate. The
measurements were 536-322-34-En 30. His stomach contained fruit and
Marmosa robinsoni chapman Allen
Didelphis (Miscoureus) murina Allen and Chapman, Bull. Amer. Mus.
Nat. Hist., V, 1893.
Marmosa marina Allen and Chapman, Bull. Amer. Mus. Nat. Hist., IX,
.Marmosa chapman Allen, Bull. Amer. Mus. Nat. Hist., XIII, 1900.
One immnature male was trapped on 31 July 1959 at the base of a
large clump of bamboo at Pointe-a-Pierre. His measurements were 1921-
7-Larmojsa carr I (Alen and Chapmsn)
E~nylamys carrt Alilen and Chaepmen, 6Eull. Am~er. Muh Nast. Hi~t., EX,
PMarosa card In Trouessert C~at. Hammn. viv. fors. 3Suppl. 856.
Aloustte seniculuc- Incularis Elliot
Myeter. g ':Enicl' Is (Linn.r, Aillen aned Chaspman, bull. Amer. Mulrs. Un~t.
Alouette seniculus insularis Elliot, Ann. and Mag~. Nat. Hist. Ser.
Hovler mocrkeys were often heard in the late afternOon Or before
rain at iuayaguayare. Ene, wre usually In bands~ of ) to 10i, each band
aFpparing to have a flxecr route t~hrourgh~ the f'orest. We observed a small
band, composed of several adults with at least oine coulnc of' that reason,
feeding on More flowers ar. Quayaguayrer on 26 AUhg)Et 1j5'. i'00 yOllcg
male of' this band was shot. His meas~ureents rere 1216-651-15-Enn y)..
Hog- plum jreeds were contrrelnred In nts stromach~ and Intestine. TNo Eskulls
were folund on th~e trail to La Table in Ganyaguaynree on t6 Au.gust 195),'.
Cebus alb~fronj trinitatis FunCh
reba sp., Thomas Trinidad Flpiel Ijt. Club I, 1893.
Cebui: s aglia, Vce~~Fr~ey-itge d (nec. Lonaeu~s!, T'ropical Ag;r.
ICtEbu albifrons trinitat~s Palson, Plejttach foir Sauget, lo, 1081.
Caspue-nni mooneys move in large bands~, 15 or more. Thier ull come
quite~ close to investigate a cell. One band attracted In EUCh & 98/
uas seen In Guaeyaguafaere forest. i'tE uale spFecimen, fDorme:rl a pet,
vas obtained from the Trinidad Regional Virus Laboratory. Its measure-
ments were 810-404-123-En 35.
Tamandua longicaudata longicaudata Wagner
Tamandua tetradactylus (Linn.), Allen and Chapman, Bull. Amer. Mus.
Nat. Hist., V, 1893.
Tamandua longicaudata Wagner, Vesey-Fitzgerald, Tropical Agr.
(Trinidad), 13(6), 1936.
One ant eater or "mataperro" was shot at Guayanguayare on 27 Aug-
ust 1959. A partial skeleton was found on the trail to La Table on 26
Dyclopes didactylus didactylus (Linn.)
Cyclothurus didactylus (Linned, in Allen and Chapman, Bull. Amer.
Mus. Nat. Hist., V, 1910.
Cyclopes didactylus didactylus (Linn.) in Vesey-Fitzgerald, Tropical
Agr. (Trinidad), 13(6), 1936.
Dasypus novemainctus novemcinctus Linn.
Tatusia novemcinctus (Linn.) in Allen and Chapman, Bull. Amer. Mus.
Nat. Hist., V, 1910.
The armadillo or "tatou" is widely distributed and common. It is
the most common large mammal in the Mora forest of southeastern Trinidad
according to hunters. One male was shot on 31 August 1959 in Gruayaguay-
are with measurements of 813-385-91-Eh 30. Three more individuals were
shot in the same area on 27 August 1959. An immature female was run
down in the cocoa at Spring Hill Estate, 13 August 1959, with measure-
ments of 670-345-80-En 37.
ciurur granatenris chapmant Alen
Mu~s. Nat. tMEt., V, 1jlO.
Edlurus chapmasnt Alen, Bull1. Amecr. Mlur. Ijt. HIpc., XIZ, lIri'D.
Thiese are fairly corimon.
Nlecrtoml; rquampi es palmpes A~llenr and Chapran
Holochilus squamipes (brhnts) In Phoase, Jour. Trinidad Firld N~at.
N:ectomyis palmliPe Allen and Chapmahn, Bull. Am~er. Mur. Ijst. Hist.,
Th~ele are commonly found at thie edge of ponds or streams~. Nvo
were trspped at the edge of a rhadlow pond In a logged area and one
war rbot along a Itream In Guayapanyi:are f'orest. Thie tw; naile vere
bothr in breedling condition and were caught the 29 and jl Atugus 1951i.
Ttbey maasured 4?7-~220:-51-E 23, Skf-17-hS-En 2l, respe~ctively. One
p'regnrant f'emale with three emb~r,s unich avera,-ed 27 IIm. crownr-rumrp use
carught the 31 Aupas~lt 195s vlth meapurements of 31_-18).-45-En 3.
Orizomva concolor speCIosuI Allen and C'hsapmn
u~ryzonver speciorus Alen and Chapmhn, Bull. Amer. PUs.. Nt. Hist.,
Orlourc-s trinitatts Allen sad Chapmian, Bull. Amer. Mlu;. list. Hist.,
IOne ~pregnan t male with two very early embryos was trapped at
Pointe-a-Pierre on 28J July 195 .:. Its measurements were 261l-136-33.
Oryzomys laticeps velutinus Allen and Chapman
Orysomy velutinus Allen and Chapman, Bull. Amer. Mus. Nat. Hist.,
Three were trapped between 13 and 16 August 1959 at Spring Hill
Estate between the cocoa and the uncleared land. There were some gnawed
cocoa pods on the ground. One female was a subadult, one was in breed-
ing condition, and one was pregnant with six embryos of which one was
resorbing. The measurements were 163-75-24-En 16, 210-109-27-En 18,
207- injured tail 87-28-En 20, respectively.
Orysomys (011goryzomys) delicatus Allen and Chapman
Orysomys delicatus Allen and Chapman, Bull. Amer. Mus. Nat. Hist.,
Twelve skulls and 52 rami of this species were found in owl
pellet material. This form is rare in collections.
Rhipidomys couesi (Allen and Chapman)
Tylomys couesii Allen and Chapman, Bull. Amer. Nus. Nat. Hist., V,
Rhipidomys couest (Allen and Chapman), Bull. Amer. Mus. N~at. Hist.,
Bygodontomys brevicauda brevicauda (Allen and Chapman)
Orysomys brevicauda Allen and Chapman, Bull. Amer. Mus. Nat. Hist.,
Akodon frustrator Allen and Chapman, Bull. Amner. Mus. Nat. Hist.,
Forty-three skulls are repreitnted in tne oul pellet matertlr.
This form Ir- commonl trapped.
A~kodion url-ht Acllen and Chapman
Abro~thrix: calieinosus (Tomues), p~rovisional reference in Allen and
Akode~cn urichi Allen and Chapman, BJull. Amer. Mus.I. Unat. Hist., LX,
31gnedon ef. hiriatus
P~enty-one skullls referable to rfigmodon are represented in the
owl pellet material. Aill thre owvl pellets Ertuied were collectted by the
late F. V. Urich at St. A~lugustne, to the best of everyone' knowoledje.
St. Augustine is located eignt miles east of Fort-oif-?pain and the Gullf
of Pariah and 35 miles~ from tnre oearest potat on the mainland. Bo~th the
numbers of animals represented and the distance froml the maiDland would
suggest that, if the 091 pellets uere indeed collected at Tt. Atugutinre,
these rate mllst have been caught o~n Prinidad and as such represent a new
record. Only minor dlifferencer uere noted unen they werej campredB~ with
fo~ur skulls8 referred to 3igmodon hirelltus.. Phree of these figodon
hin~utus: came from CaracaP and one from Ra~nch Grarnde unich Lr the
closest record of this species to Trinidad. (igacngs, chactraterie by
gorooed upper incitors, na:- a rang-e nearest to Trinidad. Ilershtocvitz
(1955~) zuglests that probably all species of Scigodon and ;Ligmove are
synonymou~s withHoon hlT~iO ispiidus3. This materil~ tends to ;Ilpport the
Idea that at least the E~ls fr00 Trinidad, the fouir sculls stu~died
from Venezulel, and rthe Ekdils Jtudied Of Signodon bi6pidus frOm Florida
are very close. Some measurements taken on the skulls from Trinidad
are given in Table 7.
Heteromyrs anomalus anomalus (Thompson), in Thomas,Trinidad Field Nat.
Club I, 1892?.
These are commonly trapped.
Rattus rattus (Linn.)
Five were trapped in areas under cultivation, inhabited areas,
and in the deep forest. One was from Pointe-a-Pierre, two were from
Spring Hill Estate, and two were from Guayaguayare.
Rattus nor-vegious (Berkenhout)
Mus musculus Waterhouse
Coendu prehensilis (Linn.)
Synetheres prehensilis (Linn.), in Allen and Chapman, Bull. Amer.
Mus. Nat. Hist., V, 1893.
A skeleton was obtained from a specimen killed about 10 August
1959 in Guayaguayare and a jaw of another was found. Another skeleton
was obtained from a hunter from Biche.
M mut pc (Linn.)
Coelogenys paca (Linna.), in Allen and Chapman, Bull. Amer. Mus. Nat.
Hist., Vr, 1893.
Paca or "Lappe" is a delicacy and as a result have become very
rare. It and the peccary are probably the most difficult game animal to
obtain in Trinidad. A skeleton of one was procured from a hunter from
WEAL-UREMEIJTL (DJ M`ILLDIMETERS) OF SIGMOrDiEN CF. HIR-'UTUI
M~easurement lilMbedr Mean R~ng~e
Ekull length 1 j6-6;
Cygornaic bireadttr j 220.0 1i. 5-33.6
Interorbital breadth 18 5.3 55- 6.i
Length of' palate 16 6.9 6. 3- 7-77
Alveolar lng~tj of
u~pp3r molar rov 16 6.3 5.S- 6-9
Length of incisive
cormotn 10 7.5 6.6- 83.
breadth of rosetrum Ilb. .-11
Dasyprocta aguti (Lian.)
Dasyp~racta rubrata Thomas, Ann. and Mlag. Nat. Hist. (7)II, 1898.
Dasyprocta aguti (Linn.), in Allen and Chapman, Bull. Amer. Mus,
Nat. Hist., V, 1893.
In Guaryaguayare the "guti" is thought by hunters to be next in
abundance after the armadillo. One was shot at Guayaguayare on 20
August 1959 One was obtained from Biche.
Fghlaggg armatus castaneous (Allen and Chapman)
Loncheres guganae Thomas, Trinidad Field Nat. Club ID7), 1892.
Loncheres castaneus Allen and Chapman, Bull. Amer. Mus. Nat. Hist.,
Echimys armatus Geoff.
Proechingn~s gayannensis trinitatis (Allen and Chapman)
E'ghingrs trinitatis Allen and Chapman, Bull. Amer. Mus. Nat. Hist.,
The "Pilori" inhabit the stream banks. One male in breeding
condition was shot in Guayaguayare on 29 August 1959. Its measurements
were 478-212-56-En1 30. One partial skull was found at the edge of the
Procyou cancrivorus cancrivorus (Cuvier) in Thomas, Trinidad Field Nat.
Club I(7), 1892.
Eira barbara trinit~atis (Thomas)
Galictis barbara (Linn. ) in Allen and Chapman, Bull. Amer. Mus. Nat.
Hist., V, 1893.
Tayra barbara trinitatis Thomas, Ann. and Mag. Nat. Hist. (7) V, 1900.
Lutra enudrir enuacrie (Cuvier)
Lutra inealaris Thomas, Ann. and Mag. Nlat. Hist. (8) 1, 1908b
Hereastes a~urpunctatuss aulrounctatus
Felic pardslie Linn.
Felis tigrina Elxl., EJlen and Ch~apcman, Bull. Amer. Mas. Flat.
Hist., EX, 1997
LEopajrdusB pardhall LIan., In Vlesey-Fitzgerald, 'Tropiedl Agr.
(Prinidad), 11(6), 1936.
Pecari tajacu (Linn.)
Dicotllea tafacu (L~inn.), in Allen and Cnapman, Bull.. Aber. Musa.
nat. Hist., V, 1893.
Esvassu Pecart F-ischer
Dicotylesi laciatus Cuvsier, in Acllen and Chapman, bull. Amer. Mue.
Nlat. Hist., V, 189].
Mazama~ americana trinitatie (A~llen)
Carisccls (Coassus) nemorivague (Cu1rier) in Aillen and Chapman~, Bull.
Amler. Flua. Nat. Hist. V, 1893.
Hbazama rur' (Cuvier)
razama trinitatie Allen, Bull. Amer. I023L. Flt. Hist., KXXiV,' 1915-
Dleer are less abundant than Dasyprocta and must be norted in-
creasin~g ly faer away from the p~opulatio~n center, Po~rt-of'-'pain kney
have one or two young, dropping them mainly in the sunmmer. ile hunter
kiiLed a deer with~ a well formed fetus in the middle of Augusat.. The
sbull and leg of one specimen was obtained unich~ was killed duaring the
spring of 195'! in "'parrk ForeEt FcereVe. A nunting party uttn dogs
killed a lactating doe with a single corpus luteum n the ovary at Taba-
quit on 26 June 1959. One buck was shot on 23 August 1959 at Guaya-
guayare that dressed out at 35 pounds. One deer was seen in the Guay-
Trinidad is part of the savannah region of South America as
defined by Cabrera (1940). In addition to savannah, however, the
region also includes the forested coastal range which is the mountain-
ous extension of the Andes. Trinidad is included in this forested see-
tion, and may be considered as the extreme northeastern end of the
coastal range. Th~is geographic character is reflected in the present
mammal fauna which is composed to a large extent (38 percent) of arboreal
forms. The savannah faunas which existed on the island in the Pleis-
tocene is represented by only one relict Form, Oryzomrys delicatus, in
the present fauna, and by one other, Odocoileus graotis, of the pre-
Westermann (1953) has stated that "The flora and fauna of
Trinidad and Tobago have strong affinities to those of the neighboring
South American continent, but quite a number of species common to the
Guianas are absent here. Their absence is due to natural circumstances
and not to extermination by man. When the present land mammals of
Trinidad are compared with the mammals of the Guiana Region and Rancho
Grande, Venezuela, as reported by Tate (1939, 1947), and those of
Northern Colombia as reported by Hershkovitz (1947, 1948, 1949, 1960),
it is found that more than twice as many (28 percent) have affinities
to the vest of Trinidad in Venezruele, Colombia, and the Andes tnan~ have
affrinities to the south in the Guriana region. Tnose f'orms with allies
to the vest are: Cebas albitroos, jlouatta senicullus, SciLrue grana-
t~ensis, iCryzomys delicatusf, Enipidomyaa couest, Akodon urichi, Heterous
ano~rualus, and thatnosa car [, whereas those With allies to the south are
Orlyzomye concolor, Lchtra enuaris, and C'alalromys philander. The re-
makingo forms (E,9 percent) haIVE relatively extensive ranLges. ;This
association~ again reflects the geographic tFes unich Trinidad Has had
with Venreuela and its recent erlological relationship with tne forested
areas. It alEo suggests that the iOriooco River CODptituteB 85 important
barrier to many videlyr ranging mamrmal rspedes.
Since a large part of this study is based on animal bones as-
sociated with archeological remains it is appropriate that some of the
problems involved in the use of such materials for zoological interpre-
tation be discussed. It is important to understand the nature of the
deposit in which the bones are found, the method of sampling, and the
cultural background of the human population represented. The nature
of the deposit may be ceremonial where animals are buried as part of a
ritual, but more often animal remains are associated with kitchen middens.
The number of different forms represented in a midden may often be cor-
related with the economy of the aborigines that made the midden. Usu-
ally gathering, hunting, and fishing peoples will depend on a greater
variety of animal foods than agricultural people. Middens are sampled
by the archeologist either randomly by a surface collection that may
be preliminary to an excavation or by an excavation that systematically
samples all parts of the midden. Each level, the smallest unit of the
sample, is of known volume and all that is found in each level is kept
separate for analysis. The zooarcheologist bases his analysis on material
sampled in the same manner.
The archeologist's goal is to reconstruct the life of the abori-
ginal group he is studying whereas the zooarcheologist's objective is to
reconstruct as far as possible the faunal characteristic of that period.
Such data contribute to the archeologist's interpretation of the rela-
tionship of the human population to its natural environment. The
analysis of particular faunal assocciations~ must rest upon a foundation
of accurate e ident ifI cat ion For this purpc~se as large series of sehLe-
tons as pojssile must be obtained and prepared. Usually between 25 per-
cent sad 75 percent of the material will be too fragBTetkry to identify.
From the identifiable portion, the relative number of individuals of
each species may be determined by calculaltingj the minimum number of
Individuals. This must be done to give equ~al consideration to each form
andg not favor those with more bones. With these data a better picture
of' the former ecology and various changes in the fauna nay be revealed.
Certain changes in the fauna within one site may become apparent when
the materials are analyzed stratigrapthlealllyor between the fauna of
the rite and the more distant past or present fauna of the same area.
Varkr with the individual species ruany reveal new forms, certain varia-
tions of recognized forms, or estealogica-l properties of a speclea or
genus. This material may provide much valuable inf'omaution unen ana-
lyzed with its origin in mind.
In this studyr of samples of past and present mannalian faunas of
Trinidad an effort was made to determine the composition of tne faunas
and to gsai some insight into the posabnle factors that might have
played a role in the formation of the faunas. It is obvious that the
fauna vill be composed of such forms as have had geographic access to
the area and whose habitat are represented. The fossil fauna existed
at a time when what is now TRinidlad was connected to thre mainland, and
savanznahs covered most of the area. Forest grew at nighier elevations
and along streams. The known fossil fauna reflects these conditions in
that it is composed principally of large herbivores of grassland
habitats. The geological events that resulted in the isolation of
Trinidad as an island also resulted in the rejuvenation of the soils
and thereby the spread of forest vegetation. Isolated in this habitat
is a recent fauna of entirely different nature. As many as 38 percent
of these mammals are arboreal and a great many more are forest dwell-
As would be expected, accompanying the changes in the composi-
tion and isolation of the faunas are changes in its affinities and per-
centage of endemism. The fossil fauna vas composed largely of species
with very wide ranges. There is no endemism among the fossil species
(vith the possible exception of Zygodontomys) since there was no iso-
lation in the uninterrupted expanse of savannah that spread across
Venezuela and Trinidad. The Recent fauna is composed to a large ex-
tent of widespread forms that originated principally from the vest of
Trinidad. Evidently the Orinoco River presented a barrier to the move-
ment of certain species from the south to Trinidad. Dule to the gradual
separation of the island from the mainland, species became isolated.
This is reflected in the extent of endemism now found in Trinidad. Of
the 29 land mammals now known from Trinidad, two Marmosa carri, and Ake-
don urichi, are endemic. Twelve endemic subspecies are also recognized.
Since the colonization of Trinidad by Europeans the isolation of
the island fauna has in one sense broken down and as a result of man's
activities the composition of the fauna has been altered. At the
present time the forest forms are decreasing in numbers as the forest
habitat is replaced by agricultural and industrial areas. With the
increase in the h~uman population from about d35,853 in 1550 to 825,700.
in 1960~ a drastic change has been seen in the abundiance1 of gamep animald.
Ihis situation is Farticularly ceriou3 because of three principal fac-
tors. T~he first of' these Le unoeoployment resulting f'rom the increase
in p'cpul~ation. The second is the good market for vild meats that
.stimulhats rhunting~ by the people. Finally the ,-ame lave, althougoe
reasonably strict, are poorly enforced. Hunters nboat wi~thout licenses,
shoot protected animals, and animals oult of season wirthout hesitation..
The clearlnc of land provides habitats f~or rodients and particularly
Rasttule and Mu~a at the explense of the native forms. A fewl of1 the native
forms such as Zyg~Oodoctomys adapted to habitati s resultirrg from~i cleared
land have been able to compete esuccesssfully with the introduced forms.
However, the persecution of game species aculd appear inevitably tc
lead to taelf extermrination if no reservoirs are provided. Perhaps the
most practical resolution would be to set aside sanctuarlee triat could
be msanaed and stringently protected for the replenishment of game in
other forested areas. The one area unich is absolutely protected nov
is the Ulnited States baee at Chagaracius where gamje is said to be Fpolet-
full. As Westeranlrn (19353, p. 123) has saidi, "The relatively EGIF11 SIZE Of
these islands, and their rapidly growlag population, ruake the oultlookh for
the preservation of nature rather precarious, unless large forest and
natulre reserves can te ktept in p~erpetuity."'
SUMMLARY AND CONCLUSIONS
Three samples of mamrmal bones representing different time peri-
ods were studied to see what changes have taken place since the late-
Pleistocene and to determine the cause of these changes.
The first of these samples was deposited more than 34,000 years
ago in a stream edged by gallery forest when Trinidad and Venezuela
were broadly connected. The mammal fauna represented was composed of
large herbivorous forms as Cuvieronius, Glyptodon, Pbylodon, and Mega-
therium of which all are now extinct. In addition to these large mam-
mals one small rodent referable to the genus Zygodoctomys was repre-
sented. Although a species of Zygodontomys now exists on Trinidad it
is not the same species as the fossil form. The fossil Zygodontomys
is characterized by its very small size. The ecological situation
indicated by the fossil mammals is that the area's habitat was grass-
land and forest.
The second sample was composed of bones excavated at six Indian
middens. The mammals represented canprise almost all the larger land
mammals native to Trinidad. Those larger than the cricetid rodent
Rhipidomys omasi that were not found are Cyclopes didactylus, Cebus al-
bifrons, and Eira barbar, although the latter has previously been re-
ported from a midden. Species that do not now occur on the island are
Odocoileus gymnortis, which probably constitutes a relict; Tagassu
pecari, which is probably now extinct on the island but did appear in
the early lists of Trinidad mammals; and Tapirus represented by one
tooth which may have been a trade object.
Tne principal difference between sites was not found to b~e the
presence or absence of' certain animals, but rather trhe percentage of
comparable and abundant forms. The armadillo was found to be more
abundant at sites where the soils are more r'riable. At the three
sites situated on the south coast in open forest, deer and agou~ti were
more abundant than FCenaTry and p8ac, respctively. At thle site in palm
forest jirrounded b~y heavy forest agouti and paca vere almost equally
abundant but deer were more abundant than pec~cary. Phe two attes
located in neavy forest had far more paca and p~ccary tnan agou~tt and
deer, respectively. These differences in the percentage of vasriouls
forms reveal more subtle differences to the ecclorr, of' an area than
are revealed b, the presence or absence o~f an incidental form.
The last sample is composed of' specimena that uere collected by
us, and by nmamml remained from owl pellets, and includes a compilation
of publiabed records. iFbe only inceported form f'ound is a rodent refer-
able to? S`ignedonr ef. Mlrsutus-. Thnis record Is based on 21 skaile from
owl p~eilles collected originally by F.'d. Urica. The R~ecent fauna is
composed to a largle extent of forest forms that are allied principally
to species from the uest of' Prinidad or *ith videspread rangjes. The~se
affinities with the west reflect tne past land connection of Trtnidad
and Venezuela and the barrier of the C'rinoco River to the disFPersal of
certain forms. Today with the removal of' forest by mao many of' the
nat ive mammal populat ions are duludl ing.
Allen, J. A. and F. M. Chapman, 1893. On a collection of mammals from
the Island of Trinidad, with descriptions of new species. Bull.
Amer. Mus. Nat. Hist., 5(13):203-234.
.1897. On a second collection of mammals from the Island of
Trinidad, with description of new species, and a note on some
mammals from the Island of Dominica, W. I. Bull. Amer. Mus. Nat.
Beard, J. S. 1916. The natural vegetation of Trinidad. Oxford Fores-
try Memoirs No. 20, pp. 152.
Blair, K. G. 1927. Remains of insects from oil sand in Trinidad. Trans.
Entom. Soc. London. 137-142.
Bullbrook, J. A. 1953. On the excavation of a shall mound at Palo Seco,
Trinidad, B. V. I. Yale Univ. Publ. Anthr. No. 50:5-114.
Burt, W. H. 1961. A fauna from an Indian site near Redington, Arizona.
J. Mamm. 42(1): 115-116.
Cabrera, A. 1957. Catalogo de los mamiferos de America del Sur. Miuseo
Argentino de Ciencias Naturales "Bernardina Rivadavia."
Cabrera, A. and J. Yepes. 1940. Mamiferos Sud: Americanos. Campania
Argentina de Edit'ores, Argentina, pp. 370.
Cruxent, J. M. and I. Rouse. 1959. An archeological chronology of
Venezuela. Vol. I and 2:1-223. Social Science Monograph 6 of
Pan American Union.
Eniliani, C. 1955. Pleistocene Temperatures. Jour. of Gee., 63(6):
Goodwin, G. G. and A. M. Greechall. 1961. A review of the bats of
Trinidad and Tobago. Descriptions Rabies infection, and ecology.
Bull. Amer. ~Mus. Nat. Hist., 122(3):191-301.
Greenhall, A. M. 1956. Is the Yapok or water opossum found in Trinidad?
Jour. Trinidad Field Naturalist's Club, p. 27.
Gyldenstolpe, N. 1932. A manual of Neotropical Sigmodont rodents.
Kungl. Svenska Vetenskapsakademiens Handlingar. 1()114
Herahkovit-, P. 196l7, blammalE Of INrthprn~ Colombia preliminary report
Ilo. 1: -.quirrels (3ciuridae). Pro~c. U. 3 lt u.9(.0).6
194:8. Mammals of Nlortaern C'olombias preliminary report Ilo. 2:
,piny rate (Schivildae), with supplemental notes on related
forms. Proc. U. S. Nlat. Mus. P7(3216):L25-140.
.1948. Mammanls of Northern Colombia preliminary report Ilo. 3:
v;ater rats (genus Ilectomysl) wlith supplemental notes on related
forms. Proc. U. ,. NJet. Mus. 98(3221):49-56.
.1949. M~aommls of Northern Colombia preliminary report No. 4:
monkeys (Prima~tes), with taxonomic revision of sjome forms. Proc.
U. s. Nat. Mus. j98(3232):323-427.
.1955. South Amritrcan marsh rats genus Holochilus vith a
-onary of sigmodout rodents. Fieldiana: Z~ool~. 37:639-673.
.1960. Mlammal of Northern Colomnbia preliminary report No. ij:
Arboreal rice rate, a systematic revision of the sulbgenusl utcongs,
genus i~ryzomyo. Proc. U. s. Nat. Mus. 110(3420):5-1:-:i68.
Ko~ldesijn, B. W. 1958. Sediments of the Paria-Trinidad shelf. Moulton
and Co.. 'c-Graveanrage, the Hague, pp. 103.
Nota, D. J. G. 1956. tediments of rth western Guiana shelf. Meldedelim-
gen van dC Land3bouunogesChool Te Wr*eningen. Ne~derland, pp. 8.jg
Osborn, H. F. 1?36. PrcbEcidea. A monograp~h of the discovery, evolu-
tion, migration and extinction of the measjodoots and elephasnts
of the orld. Amer. M~can Nat Hist., 1:1-90j2.
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Excavation at Palo Seco, PRinided. Yale Univ. Ribl. AnTnr. No.
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Senveizerieshch Palaeoonto~login ne Gelellachaftr. 5:-
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Bul. Armer. Musi. Nat. Blat. 112(2): 125-190.
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Run~. Nat. Hist. 7()112Q
.1947. A list of the mammals collected at Rancho Grande,
in a montane cloud forest of northern Venezuela. Loologica
Taylor, W. W. (ed.) 1957. The identification of non-artifactual
archaeological materials. Nat. Acad. Sci.-Hat. Res. Council.
Thomas, O. 1893. A preliminary list of the mammals of Trinidad.
Jour. Trinidad Field Naturalists' Club 1(7):158-169.
Verteuil, L. A. A. de. 1884. Trinidad: its geography, natural re-
sources, administration, present condition, and prospects.
Cassel and Co., Ltd. London, pp. 484.
Vesey-Fitzgerald, D. 1936. Trinidad mammals. Tropical Agriculture
Westermann, J. H. 1953. Nature preservation in the Caribbean. Publ.
of the Found. for scientific research in Surinam and the
Netherlands Antilles. No. 9, pp. 106.
A~nnez Elizabeth tchvarz *se born in Cambridge. M~assa-husetts,,
Ahrch :, 197.2. fhe attended rte 'bsor SichoL in BGBtGD, M836&Chlleetts,
and was graduaedp in 1'j51. In 195S abe receivcd hier beonelor ojf Arts
degree fromn Mouln t Hlcjcjle Colleg~e in Southn Haaley, Maessahu~sctts, anrd
in 1957 she r.eived her Master of Eatence degree rmn the G~olversity
During her graduate e stdies she has held gradiiate anilstantchipe
In the repartrient of Biolor,' andl Florida t~a.e Murseum. In the summer
of 1959: she? was awarded a Nahtional Scie(nce Fo)UndatlCOn .Ilnner FeCllowjhFip
for G~raduae TeaChing Assistants Jhe la presently employed as a re-
searl h asso~Ciate of the FloridaB 3~te Museulm o~n a ,ocarscheo~logcalC
project supprrted ty, the National ';ciecer Foundjation,, Grant17rr
'he is a member of` 31iga Xt and Phi 5igma ceocietesl~ an thre
American Scieti of Mamma~jlogists
in April Id, 1957, she was married to Jame~s E. Wing, Jr. and
has one chld~, Mary Elizabeth, b~nor Leptemaber 12, 191.
II cC~. .1
This j1sse~rtati~on wee prepared under the direction of the
chairman of~ the candidate's supervisory committee and has treen ap-
provcd tof all members ofP the committee. It asb submitted to the dean
o~f the College of Arts an.$ cdences and to the 3radu~ate Councl and
wee approved as partial fulfilllment of the requ~irements for the degree
of' Coctor~l of l110ophy.
ftsa. Colltege of Arts and feL~nces
tran, Grasduate Lehoocl