Timm, R. M. 1994. The mammal
auna Pp. 229-237
+
394-398
in La Sel a: Ecology and
na u al his o y o a Neo opical
ain o es (L.A. McDade, K. S.
Bawa, H. A. Hespenheide, and G.
S. Ha sho n, eds.). Uni e si y
......&
"'1L..! _ _
U.1.
ll.l.cago. ' ess,Chicago, 486
pp.
The Mammal Fauna
Robe M. Ti n n
La Sel a has been a ield s a ion o he O ganiza ion o T opi-
cal S udies (OTS) mo e han a qua e -cen u y. Habi a and
wildli e ha e been p o ec ed o managed since he mid-1950s
when Leslie R. Hold idge c ea ed Finca La Sel a. In his
chap e I explo e he na u e and di e si y o he mammalian
auna o La Sel a by in eg a ing wha is known abou his
auna om he published li e a u e in he con ex o my own
imp essions and expe ience. I p esen a comple e lis o he
mammals ha a e known o occu a La Sel a wi h es ima es
o hei abundance in appendix 8 and discuss he e he dis ibu-
ional pa e ns o hese species. I also discuss esea ch pub-
lished on mammals a La Sel a and b ie ly e iew some o
he mo e signi ican esea ch indings. I compa e his auna
b ie ly wi h o he opical aunas, discuss he impac o human
ac i i ies on mammalian dis ibu ions and abundance in his
egion, and no e some o he majo gaps in knowledge as sug-
ges ions o u u e esea ch.
THE MAMMALIAN FAUNA OF COSTA RICA AND
LA SELVA
Cos a Rica is one o he ew coun ies in he Wes e n Hemi-
sphe e in which he en i e mammalian auna ha was p esen
a he ime o Eu opean se lemen is s ill la gely ex an . Good
popula ions o mos species can be ound wi hin he coun y:
popula ions o a ew species, howe e , ha e been educed o
sizes ha can no longe be conside ed heal hy and iable. O
he app oxima ely 116 species o mammals o iginally ound
a La Sel a (app. 8), pe haps only wo species, he gian an -
ea e and he whi e-lipped pecca y, ha e been locally ex i -
pa ed. Bo h species may ha e disappea ed om La Sel a since
he 1970s (Timm e al. 1989). Al hough occasionally p esen ,
jagua s a e now only a ely obse ed a La Sel a and do no
ha e a iable b eeding popula ion he e. Thus, La Sel a e-
ains mo e han 98% o i s componen species.
The mammalian auna o La Sel a his o ically included 5
ma supials, 65+ba s, 3 (possibly 4) p ima es, 7 eden a es, 1
abbi , 3 squi els, a leas 9 long- ailed a s and mice ( amilies
He e omyidae, Mu idae, and Echimyidae). 1pocke gophe . I
po cupine, I paca, 1 agou i, 5 mus elids, 4 p ocyonids. 5 ca s.
2 pecca ies, 2 dee . and 1 api (app. 8). These 116 species
ep esen 57% o he species o mammals ound in he coun y
(app. 8). This lis canno be conside ed a comple e enume a-
ion o all mammal species ha ha e occu ed o cu en ly do
occu a La Sel a, howe e . Two species o ba s, P e ono us
da yi and Ch o op e us au i us, ecen ly we e added o he
aunal lis and, undoub edly, addi ional species o ba s and o-
den s will be ound he e (Timm e a!' 1989). Wi h he excep-
ion o hese o de s i is likely ha he lis o La Sel a's mam-
mals is comple e.
This lis o La Sel a mammals di e s om ha o Wilson
(1983) and unpublished lis s ha ha e been ci cula ed o e he
yea s in ha some species ha e been added and se e al spe-
cies now suspec ed no o occu he e ha e been dele ed. Iden-
i ica ions o specimens in collec ions ha e been e i ied o
co ec ed, and all published li e a u e eco ds ha e been e al-
ua ed. No species a e included as "expec ed o occu in he
a ea," a ca ego y ha c ea ed an a i icially high species coun .
I base es ima es o ela i e abundance o species on my obse -
a ions and hose o o he expe ienced indi iduals and on he
logbook o mammal sigh ings kep a La Sel a since 1979,
which I c i ically e alua ed. The 116 documen ed species a e
21 ewe han Wilson's (1983) lis ing o 137 species known o
expec ed o occu a La Sel a: i is unlikely ha hese "ex-
pec ed" addi ional species occu in he Pue o Viejo egion.
The majo i y o mammals ound a La Sel a a e wide-
sp ead species. ypical o neo- opical ain o es s (app. 8).
Mos ha e a b oad dis ibu ion h oughou he opical low-
lands o Cen al Ame ica. Many species a e e en mo e widely
dis ibu ed. including some wi h b oad ele a ional anges in
Cos a Rica (and elsewhe e in Cen al Ame ica) and/o wide
geog aphic dis ibu ions h oughou he lowlands o Cen al
and Sou h Ame ica. O he 116 species o mammals ha oc-
cu ed a La Sel a, 58 (49%) a e b oadly dis ibu ed in he
Neo opics, o en occu ing as a no h as Mexico and ang-
ing sou h h ough Cen al Ame ica and much o opical Sou h
Ame ica: 35 (30%) occu in he no he n opics, gene ally
being ound om opical Mexico h ough Cen al Ame ica
and no he n Sou h Ame ica; 15 (13%) a e es ic ed o Cen-
al Ame ica; 7 (6%) a e e y wide anging, ound om he
Uni ed S a es h ough Cen al Ame ica and ac oss mos o
Sou h Ame ica; and 1 (1
o/c)
is widely dis ibu ed in No h
Ame ica, occu ing as a sou h as cen al Panama (app. 8).
In e es ingly, he ba auna o La Sel a consis s o e whelm-
ingly o species ha a e ei he widely dis ibu ed in he opics
o a e o no he n Neo opical dis ibu ion (60 o 65 species,
o
92o/c).
La Sel a's p ima es a e all o no he n Neo opical
dis ibu ion. The eden a es a e all o ei he wide dis ibu ion in
he opics o o no he n Neo opical dis ibu ion. The single
species o abbi ound a La Sel a has a wide- anging dis i-
230 ROBERT M. TIMM
bu ion in he lowland opics o Cen al and Sou h Ame ica.
The oden auna a ies: some membe s a e widesp ead
in he opics, whe eas o he s a e o no he n Neo opical o
Mesoame ican dis ibu ion. All o he smalle oden s, includ-
ing he squi els, pocke gophe , long- ailed a s and mice, and
he po cupine, a e ei he o no he n opical o Mesoame ican
dis ibu ion, whe eas agou is and pacas a e widely dis ibu ed
in he opics. The ca ni o e auna is ep esen ed by many spe-
cies whose anges include much o he opics bu also in-
cludes species wi h mo e ex ensi e non opical dis ibu ions
(7 o 15 species, o 47%).
In gene al, species ha a e common a La Sel a a e com-
mon h oughou hei anges, and species ha a e a e a La
Sel a also a e a e h oughou hei anges. No able excep ions
in he ba auna a e Ca ibbean whi e ba s, which a e mo e
abundan a La Sel a han a any o he si e om which hey
a e known and disk-winged ba s, which a e abundan a To u-
gue o and elsewhe e in he Ca ibbean lowlands bu a e a e a
La Sel a. Al hough his a i y is unexplained, i is no because
o he absence o sui able oos ing si es in he Pue o Viejo
egion. These iny insec i o ous ba s oos only in he imma-
u e, olled lea es o Heliconia, which a e abundan a La
Sel a, and in which he ba s a e easily loca ed i p esen .
Rela i ely ew species o mammals a e endemic o Cos a
Rica: six oden s, one ca ni o e, and, pe haps, wo o h ee
sh ews. Mos o hese a e species o mid- o high ele a ions
and occu in es ic ed habi a s. O hese endemics, only he
pocke gophe , O hogeomys che iei, occu s a La Sel a. O -
hogeomys che iei is ound in a b oad ele a ional band no h
and eas o he Co dille a Cen al and no h and wes o he
Co dille a de Tila an in no he n Cos a Rica (Ha ne and
Ha ne 1987). The Ca ibbean whi e ba is he only o he mam-
mal species a La Sel a ha has an ex emely limi ed dis ibu-
ion. Whi e ba s a e es ic ed o he Ca ibbean lowlands o
Hondu as, Nica agua, Cos a Rica, and ex eme no hwes e n
Panama (Timm 1982).
Th ee g oups o mammals ound elsewhe e in he Neo op-
ics a e absen om he auna o he La Sel a egion: sh ews
(So icidae), coyo es and oxes (Canidae), and bea s (U sidae).
A single genus o sh ew, C yp o is, is ound in he Neo opics,
and a leas i e species occu in Cos a Rica, bu all a e e-
s ic ed o he highlands (Woodman 1992). Canids do no
occu in he Ca ibbean lowlands o Cos a Rica al hough he
amily is widely dis ibu ed wo ldwide and occu s h oughou
No h, Cen al, and Sou h Ame ica. The bush dog, Speo hos
ena icus, does (did) occu in eas e n Panama, including he
Ca ibbean lowlands. Coyo es, Canis la ans, con inue o ex-
pand hei ange in Cos a Rica (Vaughan 1983: Monge-N. and
Mo e a 1987) bu ha e ye o each he Ca ibbean lowlands.
Bea s ha e a disjunc dis ibu ion in he New Wo ld, being
widely dis ibu ed in No h Ame ica, absen in Cen al
Ame ica, and wi h a single species, he spec acled bea
T em-
a c os oma usi, occu ing in he Andes o no he n Sou h
Ame ica.
His o ical biogeog aphy o he Cen al and Sou h Ame i-
can mammal aunas was e iewed by He shko i z (1972).
Sa age (1974, 1982). Ma shall e al. (1982), and Rich and
Rich (1983) and o Cos a Rican oden s by McPhe son (1985,
1986). An in e es ing e iew o mammalian ecology in Cos a
Rica is p o ided by Janzen and Wilson (1983): i includes in-
sigh ul discussions o he ela i ely low species di e si y
o e es ial mammals, seasonal en i onmen al s ess, he
la gely ugi o ous die o many opical "ca ni o es," and an
o e iew o opical ba adia ion. They poin ed ou ha he
high species di e si y o mammals seen pe uni a ea in Cos a
Rica and elsewhe e in Cen al Ame ica is he esul o a d a-
ma ic inc ease in he numbe o ba species. Species ichness
o e es ial mammals pe uni a ea in opical Cen al
Ame ica is simila o ha o empe a e No h Ame ica. Excel-
len summa ies o he biology o many o he common mam-
mals a La Sel a and in Cos a Rica may be ound in Cos a
Rican Na u al His o y (Janzen 1983b). Two ex emely use ul
and nicely illus a ed guides o Neo opical mammals, many
o which occu a La Sel a, we e published. Eisenbe g (1989)
ea s dis ibu ion and iden i ica ion o he mammals o he
no he n Neo opics (de ined as Panama, Colombia. Venezu-
ela, and he Guyanas). Emmons and Fee (1990) is a ield
guide o Neo opical ain o es mammals and co e s species
occu ing p ima ily below 1,000 m in ele a ion.
RESEARCH ON MAMMALS AT LA SFLVA
Un il he 1980s he no he n Ca ibbean lowlands and
midele a ional slopes o Cos a Rica ecei ed li le a en ion
om na u alis s, pa icula ly mammalogis s. Goodwin
(1946) discussed he dis ibu ion and na u al his o y o he ap-
p oxima ely 125 species o mammals hen known o hough
o occu in he coun y in his Mammals o Cos a Rica, bu he
made li le men ion o he no heas e n lowlands and ci ed no
specimens om he egion. E en oday e y ew specimens
o mammals om his egion eside in scien i ic collec ions,
which hinde s e o s o be e unde s and he auna.
I was no un il he 1960s ha s udies on mammals began
a La Sel a. The i s published men ion o mammals he e
and in he su ounding egion was by Paul Slud (1960, 76) in
his classic s udy o he a i auna. In he 1960s a ew pape s
we e published on a e species o mammals ound a La Sel a.
As OTS de eloped La Sel a in o a wo king biological s a ion,
i s popula i y wi h esea che s g ew. In he 1970s and 1980s
nume ous publica ions on a wide a ay o subjec s appea ed
on mammals a La Sel a. These s udies can be b oadly ca ego-
ized in o ou main a eas: dis ibu ion and sys ema ics. ecol-
ogy and na u al his o y, communi y s uc u e, and mammal-
plan and mammal-insec in e ac ions
Pape s ocusing on geog aphic dis ibu ions and sys ema -
ics o en include mo phome ic da a, axonomic no es, and
aluable na u al his o y in o ma ion, especially on ep oduc-
ion. Se e al dis ibu ional pape s deal exclusi ely wi h ba s
(Casebee e a!. 1963; S a e and Casebee 1968; Ga dne e
al. ]970; LaVal 1977), and one deals wi h he possible occu -
ence a La Sel a o nigh monkeys o he genus
Ao U5
(Ti nm
1988). Wilson's (1983) checklis o mammals a he OTS ield
si es was he i s mode n assessmen o mammalian dis ibu-
ions in Cos a Rica, and as such gi es esea che s an impo an
baseline. Two ecen epo s e iew he auna o his egion
(Ti nm e al, 1989; Wilson 1990).
Timm e al. (1989) conduc ed a aunal su ey o he ele a-
ional ansec om 35 m o 2,600 m ha encompasses La
Sel a and Pa que Nacional B aulio Ca illo o Volcan Ba a.
We documen ed ha a leas 141 species o mammals occu in
he egion, including se e al species new o he a ea's auna.
Addi ionally, we e iew he sys ema ics. dis ibu ion. and na -
u al his o y o hese species and p o ide new biological in o -
ma ion on each. This is he i s in-dep h s udy o an ele a-
ional ansec in he Neo opics, and he i s comp ehensi e
e iew o he mammalian auna o any egion o Cos a Rica
since Goodwin's (1946) Mammals o Cos a Rica.
Publica ions om La Sel a in he ca ego y o ecology and
na u al his o y o mammals include s udies on na u al his o y.
beha io , gene al ecology, and e olu iona y ecology. Empha-
sized he e a e speci ic ecological p oblems as well as in-dep h
au ecological s udies. In e es ingly, mos o he wen y-odd
published s udies in his ca ego y ha e been on ba s. In addi-
ion, Fleming has published se e al epo s o his s udies on
oden s; G eene (1989) p o ides in e es ing obse a ions o
agg essi e in e ac ions be ween male h ee- oed slo hs; and
Fishkind and Sussman (1987) p o ide a p elimina y su ey o
p ima e densi ies a La Sel a and he adjacen zona p o ec o a
(now Pa que Nacional B aulio Ca illo), sugges ing ha six o
en g oups o howle monkeys, wo o ou g oups o spide
monkeys, and wo o h ee g oups o whi e- aced capuchins
a e in he La Sel a ese e. Two impo an s udies, bo h on
p ima es, a e unde way a La Sel a in 1992. Amy Fishkind
Campbell is s udying whi e- aced capuchins, and Ka h yn
S one is wo king on howle monkeys. These s udies ep esen
he i s modem au ecological wo k o be unde aken on p i-
ma es in he Ca ibbean we o es s o Cos a Rica and as such
will p o ide e ined es ima es o densi ies and aluable com-
pa a i e da a.
Se e al species o ba s, including some o he mos poo ly
known and hose conside ed o be ex emely a e, ha e been
s udied a La Sel a. Un il i was edisco e ed a La Sel a in
1961, Ec ophylla alba ( ig. 18.1) was among he leas known
o all mammals: only wo poo ly p ese ed specimens exis ed.
one collec ed in he la e 1880s and om Hondu as and one in
he ea ly 1890s om Nica agua (see Timm e al. 1989), wi h
no associa ed biological da a (Casebee e al. 1963; Timm
1982). Se e al aspec s o i s biology ha e been in es iga ed a
La Sel a, including i s cons uc ion o oos " en s" by modi-
ying lea es (G eenbaum e al. 1975; Timm and Mo ime
1976; LaVal and Fi ch 1977; Timm 1982; Timm and Ke mo
1982; B ooke 1987a, 1989). Whi e ba s c ea e en s by se -
e ing he la e al eins and in e connec ed issues on bo h Heli-
conia and Cala hea lea es om nea he base o he ip o he
lea , causing he sides o he lea o d oop down. The oos ing
ba s hang om he mid ibs o he lea es and a e p o ec ed
om bo h p eda o s and he elemen s by hei en s ( ig. 18.1).
Whi e ba s oos only unde hese al e ed lea es, using hem
as diu nal oos s, ma e ni y oos s, and nigh - eeding oos s.
Ec ophylla alba is now known om hi een locali ies, bu
only a La Sel a and in he adjoining Pa que Nacional B aulio
Ca illo is he e assu ance ha i and i s habi a will be p o-
ec ed. O he s udies on speci ic aspec s o he ecology o ba s
a La Sel a ocus on social o ganiza ion and o aging in em-
ballonu id ba s (B adbu y and Veh encamp 1976b), en con-
s uc ion and social o ganiza ion in Vampy essa nymphaea
(B ooke 1987b). p ey selec ion and oliage gleaning by
Mi-
c onyc e is megalo is (La al and La al 1980). ep oduc ion
du ing he d y season (Ma es and Wilson 1971), he in luence
o human-made ails on o aging by ugi o ous ba s (Pal-
THE MAMMAL FAUNA 231
Fig. 18.1. Pho og aph o a colony o ou whi e ba s. Ec ophylla alba.
oos ing in a en cu om a Heliconia lea . The ba s ypically hang cu led
in igh clus e s om he mid ib. De ails o he cu side eins and in e con-
nec ed issues may be seen along he mid ib o he lea . The holes in he
lea we e made by he ba s' claws. Ten s such as his one may be used
o se e al weeks. Ec ophylla oos s only unde lea es ha i modi ies
as en s.
mei im and E he idge 1985), he in luence o body size on die
and habi a in Ca ollia (Fleming 1991), ecology and sys ema -
ics o en -making ba s (Timm 1984, 1985, 1987; Timm and
Clauson 1990), o aging o ampi e ba s (Young 1971a), and
ecology o Saccop e yx bilinea a (Young I972c; bu see
1975).
In he ea ly 1970s Theodo e H. Fleming began a se ies o
classic s udies on he popula ion ecology o Desma es 's spiny
pocke mouse, He e omys desma es ianus (see Fleming 1973,
1983; Fleming and B own 1975; and addi ional e e ences
ci ed). He in es iga ed popula ion dynamics (Fleming 1974a),
social o ganiza ion (Fleming 1974b), g ow h and de elop-
men (Fleming 1977a), and expe imen al esponses o animals
o manipula ed ood and wa e a ailabili y (Fleming 1977b).
These s udies we e among he i s in-dep h on any species o
small opical oden . As such, hey p o ide an impo an basis
o compa isons wi h empe a e oden s and some o he i s
baseline da a on opical species. Fleming's wo k is especially
impo an in elucida ing he ecology o and oles played by
bo h H. desma es ianus and Liomys sal ini as "key indus y"
species in opical o es s. As hese he e omyids a e o en he
mos abundan small mammals in he communi y, hey se e
as bo h majo seed p eda o s and seed dispe se s and a e im-
po an p ey i ems o a wide a ie y o ca ni o es.
A La Sel a Fleming ound He e omys o be he mos
abundan small e es ial mammal p esen . Densi ies anged
om nine o eigh een pe hec a e and he popula ion was
s able o e he wo-yea s udy. Bo h males and emales b ed
h oughou he yea al hough ep oduc i e ac i i y declined
ma kedly du ing May and June. Li e size a e aged 3.1
young, and emales p oduced up o i e li e s pe yea . He e -
omys is pe haps he dominan g ani o ous e eb a e o he
o es loo . Fleming ound spiny pocke mice o eed ex en-
si ely on seeds o he palms Soc a ea du issima and Wel ia
geo gii. and we obse ed hem o eed on he palms Eu e pe
mac ospadix, Geonoma sp.. and I ia ea gigan ea and on Mel-
iosma sp. (Sabiaceae) (Ti n n e al. 1989). Based on ex ensi e
232 ROBERT M. TIMM
apping, Fleming sugges ed ha He e omys does no ha e
mu ually exclusi e home anges o e i o ies. My li e ap-
ping o his species a Mon e e de ce ainly suppo s ha con-
clusion. The e a e, howe e , emendous popula ion luc ua-
ions o spiny pocke mice a Mon e e de (pe s. obse a ion).
Publica ions ha ocus on mammalian communi y s uc-
u e a La Sel a a e exclusi ely conce ned wi h ba s (Findley
1976; Findley and Wilson 1983; LaVal and Fi ch 1977). One
o he i s compa a i e s udies o opical ba s was by LaVal
and Fi ch (1977), who compa ed he s uc u e, mo emen s
and ep oduc ion o he complex ba communi ies a La Sel a,
La Paci ica ( opical d y o es ), and Mon e e de (p emon ane
mois and p emon ane we o es s). They ound he highes
species di e si y o ba s a La Sel a; much o his di e si y
was due o insec i o ous ba s, bo h oliage-gleaning and
ae ial eede s. The h ee si es we e simila in species di e si y
o nec a - and pollen- eeding ba s. Mos opical ba s ep o-
duce seasonally. Ba s a La Pac ica. wi h i s sha ply deline-
a ed we and d y season. ha e he sho es and mos sha ply
delinea ed ep oduc i e seasons. A longe ep oduc i e season
was ypical o ba s a La Sel a, and Mon e e de was in e me-
dia e be ween he wo (LaVal and Fi ch 1977).
I is o en assumed ha he ex eme species ichness ob-
se ed in a la ge, complex auna like ha o opical ba s can
occu only i species a e es ic ed o na ow and mu ually
exclusi e eeding niches. Findley (1976), howe e . compa ed
he ba auna o La Sel a o hose a o he opical and empe -
a e si es and demons a ed ha al hough empe a e aunas do
exhibi g ea e a i y o e en absence o ecologically dis inc-
i e axa compa ed o opical aunas, opical and empe a e
aunas do no di e signi ican ly in species packing o deg ee
o niche o e lap among componen species.
Findley and Wilson (1983) demons a e ha species den-
si y o New Wo ld ugi o ous ba s is signi ican ly g ea e
han opical A ican ugi o ous ba s. Cap u e a es wi h
s anda d mis ne s showed ha he absolu e numbe s o spe-
cies cap u ed, as well as he numbe s o indi iduals wi hin
species, a e highe in he Neo opics. They did. howe e , ob-
se e a compensa o y end in biomass: he la ge ugi o ous
p e opid ba s o A ica (mean body mass o 52 g ams) oc-
cu ed in oughly he same o al biomass densi y as he mo e
nume ous bu smalle New Wo ld ugi o ous phyllos omids
a La Sel a (mean body mass o 18 g ams). I should be no ed
ha he di e ence in size be ween p e opid ba s and mic ochi-
op e ans is also a phylogene ic one.
One o he ad an ages o a hea ily used biological s a ion
such as La Sel a is ha biologis s wi h di e se in e es s and
skills a e able o exchange ideas and expe ise. Such a s imu-
la ing a mosphe e encou ages c oss-disciplina y collabo a-
ion, such as he e y ui ul esea ch on mammal-plan in e -
ac ions, including s udies on seed dispe sal by mammals
(Vande mee e al. 1979), seed hoa ding (an ex emely un-
common beha io among Neo opical mammals) by spiny
pocke mice (Vande mee 1979), he e ec o p eda ion on
seeds and seedlings by mammals (Denslow and Moe mond
1982; McHa gue and Ha sho n 1983), ba pollina ion o
lowe s (Voss e al. 1980), he ole o ba s in dispe sing Pipe
(O'Donnell 1989). and he al e a ion o lea shape by ba s o
p oduce diu nal oos ing s uc u es (Fos e and Timm 1976:
B ooke 1987a, 1987b: Timm 1987).
Two impo an s udies o mammal-insec in e ac ions ha e
been comple ed a La Sel a. An appa en ly pho e ic ela ion-
ship be ween He e omys desma es ianus and a newly disco -
e ed species o ineid mo h (Amyd ia sel inii has been e-
po ed (Da is e al. 1986). Females o his mo h we e ound
only on spiny pocke mice; he male o he species is un-
known. Amyd ia sel ini has been ound only a La Sel a al-
hough I ha e sea ched unsuccess ully o i elsewhe e in
Cos a Rica a highe ele a ions. The igh associa ion be ween
emale mo hs and He e omys sugges s ha he li e cycle o
he mo h is associa ed wi h he nes ing biology o he oden s.
bu his has ye o be demons a ed. A pa allel associa ion is
ound be ween h ee- oed slo hs and slo h mo hs. The slo hs
a La Sel a ha e la ge popula ions o hese pho e ic/ec opa a-
si ic mo hs. bu hey ha e no been s udied he e. A e iew o
he complex pa asi e auna o slo hs and i s in e es ing biolog-
ical ela ionships, based p ima ily on esea ch conduc ed in
Panama and B azil. is p o ided by Waage and Bes (1985).
Th ee species o ba lies o he amily S eblidae ha a e hos -
speci ic, blood- eeding ec opa asi es on Ca ollia pe spicilla a
we e in es iga ed a La Sel a by F i z (1983,. He ound ha
he li e cycles o he ba lies a e igh ly synch onized wi h he
li e cycles o hei hos s.
HISTORY OF KNOWLEDGE OF LA SELVA
MAMMALS
Biologis s in a wide a ay o ields ha e long been awa e o
he emendous di e si y o o ganisms ound in he opics.
Wi h he ecen awa eness o he pligh o opical ain o es s,
he e has been a pa allel scien i ic awakening and in e es in
his di e si y. To unde s and he s uc u e and na u e o opi-
cal ain o es s biologis s ha e a emp ed o cha ac e ize, com-
pa e, and con as he di e si y o pa icula o ganisms a and
be ween si es.
C i ical o hese ypes o s udies is he his o ical compo-
nen o ime and how one's knowledge o he bio a a a pa icu-
la si e inc eases wi h ime. The ime componen is qui e com-
plex and includes wo aspec s: he pe cen age o he o al
auna a ailable o sampling a he ime o he s udy and he
pe cen age o he o al auna known o he in es iga o (s). In-
comple e abula ions migh esul om local ex inc ions
be o e in es iga ion, om species ha a e a e o di icul o
cap u e o obse e, o simply om lack o esou ces o ade-
qua e sampling.
Because such an excellen eco d o he mammalian auna
a La Sel a now exis s, i is use ul o examine he a e o in-
c ease in knowledge o wha species o mammals occu a La
Sel a-a species disco e y cu e o he auna.
The ques ion I add ess he e is How long did i ake o each
he cu en le el o knowledge? and he da abase I use is he
published li e a u e. A e iew o he li e a u e on esea ch a
La Sel a con ains mo e han o y- i e p ima y e e ences
co e ing he hi y-one-yea pe iod om 1960 o 1990.
The mammalian auna o La Sel a. as ecen ly as he
1960s, consis ed o a leas 116species (app. 8). The disco e y
cu e illus a es he conside able ime i ook o acqui e his
knowledge as measu ed by published epo s in he li e a u e
( ig. 18.2). Only 16o/co he auna had been iden i ied by
1970: 84o/c
was iden i ied by 1980. I was as la e as 1986.how-
120
100
(/)
Q)
o
80
Q)
Co
(/)
'0
60
•..
Q)
~
E40
:J
Z
20
01965 1975
Yea
THE MAMMAL FAUNA 233
e e , be o e 95% o he species known o be p esen we e doc-
umen ed he e. Thus, a ull wen y-six yea s elapsed be ween
he i s epo s and documen a ion o 95% o he mamma-
lian biodi e si y.
This ime lapse a such a well-s udied si e makes i clea
ha in es iga o s need o exe cise ex eme cau ion in making
compa isons be ween si es, especially in he opics. Wi hou
a su icien da abase and his o ical pe spec i e, he compa i-
sons made could be mo e misleading han insigh ul. Fo in-
s ance, si es such as La Sel a and BCI bo h app oach ha ing
100% o he mammalian di e si y iden i ied. Mos o he Neo-
opical si es ha e been s udied much less in ensi ely and o
a much sho e ime and a much lowe pe cen age o hei au-
nas is likely known. Di ec compa isons be ween La Sel a o
BCI and hese o he si es could, hus, p oduce spu ious o mis-
leading esul s.
LA SELVA AND OTHER TROPICAL SITES
Wi h he p e ious ca ea s in mind compa isons be ween he
mammalian auna a La Sel a and o he opical si es can be
use ul in unde s anding he s uc u e and complexi y o opi-
cal ecosys ems on a b oad le el. Published aunal lis s o
mammals a Neo opical si es a e ew, and a emp s o make
in e si e compa isons o opical mammal aunas ha e been
hampe ed by lack o da a on p esence and abundance o spe-
cies (see Eisenbe g and Tho ing on 1973; Emmons 1984;
Bou lie e 1989).
Ba o Colo ado Island (BCl) in cen al Panama is he only
Neo opical si e ha has been s udied in ensi ely o a longe
pe iod han La Sel a; he i s mammalogis s isi ed he a ea
in he la e 19 lOs. La Sel a and BCI a e in close geog aphical
p oximi y and ha e conside able aunal o e lap. Ba a Colo-
ado Island was isola ed as an island when he R o Chag es
was dammed o o m he cen al wa e way o Panama Canal
in 1914. Na u alis s i s isi ed BCI in 1916. and i was de-
cla ed a ese e and opical ield s a ion in 1923. Al hough
he e a e nume ous di e ences be ween La Sel a and BCI in
clima e, geology, and he esul ing o es , onesigni ican di -
e ence now a ec ing he mammals o hese wo lowland
opical si es is he ac ha BCI has been an island o se en
1985
Fig. 18.2. Cumula i e species disco -
e y cu e o mammal species known
om he La Sel a Biological S a ion.
Da a poin s a e plo ed cumula i ely by
yea and ep esen he numbe o spe-
cies added o he auna based upon he
published li e a u e. The X axis is he
da e o publica ion o he p ima y e e -
ence o he nea es qua e yea .
decades, whe eas La Sel a lies on he mainland. Addi ionally,
BCI has no s eams ha low yea - ound.
BCI has been he si e o excellen long- e m s udies on se -
e al species o mammals, including whi e- aced capuchins,
howle monkeys, slo hs, ed- ailed squi els (Sciu us g a-
na ensis), agou is, and coa is (see Leigh e aI. 1982, and e e -
ences he ein). No s udies, howe e , ha e been published
abou he La Sel a and BCI mammal aunas ha a e di ec ly
pa allel. making ecological compa isons be ween he wo di -
icul . As wi h La Sel a, se e al. species o mammals ha e
bene i ed om he educ ion and/o elimina ion o p eda-
o s by humans. The la ge ca s and ap o s, especially ha py
eagles, a e absen o a e on BCI.
A compa ison o he numbe s o species ep esen ed in
each o de o mammals in all o Cos a Rica, a La Sel a, and
a BCI shows ha he La Sel a and BCI aunas a e ex emely
simila ( able 18.1). La Sel a and BCI ha e abou he same
numbe o ma supials ( i e and six, espec i ely), nei he has
any insec i o es, and La Sel a has mo e species o ba s han
BCI (six y-six compa ed wi h i y-six species). BCI may ha e
only one addi ional species o p ima e (c es ed ba e- aced
a na ins, Saguinus oedipus); he wo si es ha e he same num-
be o eden a es, abbi s, a iodac yls, and pe issodac yls; La
Sel a, howe e , appea s o ha e wice as many ca ni o es
( ou een compa ed o se en) and 60% mo e oden s (six een
compa ed o en). As wi h he La Sel a auna, much o he
BCI auna has a widesp ead dis ibu ion in he opics, which
accoun s o he ac ha he wo si es sha e many species.
Jus how ep esen a i e he auna and densi ies o mam-
mals cu en ly occu ing on BCI is o wha one would expec
o a p is ine lowland o es in cen al Panama has been he
subjec o much deba e. Glanz (1982, 1990) documen ed bo h
he his o ical and ecen changes in abundance o e es ial
mammals on BCI, including he ex inc ion o pumas (Felis
concolo ) and whi e-lipped pecca ies, and he appa en en-
old inc ease o agou is and squi els, among o he s. He did,
howe e , sugges ha many species migh jus appea o be
mo e abundan because hey a e less wa y now as a esul o
p o ec ion om hun ing.
The auna o La Sel a is much mo e e es ial han he
aunas ound in he g ea e Amazon Basin. Faunas o season-
234 ROBERT M. TIMM
Table 18.1 Compa ison o mammalian aunal di e si y a he o dinal
le el be ween Cos a Rica in o al. La Sel a. and Ba o Colo ado Island
O de Cos a Rica La Sel a BCI
Ma supialia 8 5 6
Insec i o a 5 0 0
Chi op e a 105 65 56'
P ima es 5 3 5
Edema a 7 7 7
Lagomo pha 3
I
1
Roden ia 45 16 10
Ca ni o a 22 14 7
A iodac yla 4 4 4
Pe issodac yla 1
I I
TOTALS 205 116 97
'Handley e aJ. 1991.
ally looded o es s such as hose o Manaus a e ypically
composed o a g ea e p opo ion o a bo eal species. Al-
hough a bo eal ma supials, p ima es, oden s, and ca ni o es
a e p esen a La Sel a, he numbe o a bo eal species in each
g oup is conside ably ewe han seen a Amazonian si es. The
pauci y o a bo eal species is no a e lec ion o he o es bu
o geog aphy; mos o he uly a bo eal species a e o sou h-
e n o igin.
Te bo gh (1988, I990b ) p o ided hough -p o oking, con-
o e sial essays on he impo ance o la ge ca ni o es in
main aining biological di e si y o bo h plan s and animals in
he Neo opics. He no ed ha BCI, an island wi h ew o i s
la ge p eda o s le , has popula ions o agou is, pacas, and
coa is ha a e en imes highe han hose obse ed a Cocha
Cashii (wi h i s p eda o popula ions in ac ) in Amazonian
Pe u and ha popula ions o smalle p ey species, such as co -
on ail abbi s, a madillos, and opossums, a e om wo o en
imes g ea e on BCl. He sugges s ha a such high densi ies,
hese species ac as signi ican seed p eda o s and ha he a i-
icially ele a ed abundances o hese mammals ha occu on
BCI may ha e had a majo impac upon he s uc u e o he
o es . Al hough in e es ing, hese so s o compa isons be-
ween si es a e la gely un es able gi en he mul i ude o ac-
o s in ol ed. Fo es ype di e s om si e o si e, and he his-
o y o land use o each o he si es is qui e di e en . Fo
example, Manaus (B azil) has no o iously poo soils and a
s ong d y season, bo h o which may con ibu e o he depau-
pe a e mammal auna and low numbe s o indi iduals he e
(Malcolm 1990). Manu (Pe u) has hi een species o p i-
ma es, many o which occupy a squi el-like niche, pe haps
con ibu ing o i s pauci y o squi els. The e is li le accep-
ance among wildli e manage s o he no ion ha a single
p eda o species can ac ually con ol o egula e a p ey spe-
cies' popula ion. Elimina ion o many o all p eda o species
coupled wi h majo habi a changes. howe e , may impac
p ey species' popula ions.
IMPACT OF HUMAN ACTIVITIES
Se e al species o mammals a La Sel a undoub edly ha e in-
c eased in abundance as human ac i i ies adjacen o he e-
se e opened he o es and as he ese e expanded o include
seconda y o es and o he dis u bed a eas. Con e sely, hun -
ing p essu e and o es des uc ion ha e a di ec nega i e
impac upon he numbe o species and, o en, densi ies o
mammals.
Subsis ence a ming. especially when based on small am-
ily ga den plo s su ounded by o es , has a posi i e e ec on
species di e si y and abundance o ce ain ma supials, ba s,
and small o medium-sized oden s. C ops and hei associ-
a ed insec pes s p o ide a dense and eadily a ailable sou ce
o ood o animals. Addi ionally, opening up o he o es in-
c eases he edge e ec , o eco one habi a , c ea ing a ich hab-
i a o many species. Species ha ha e undoub edly inc eased
in abundance because o human ac i i y in he egion include
Didelphis ma supialis, Glossophaga so icina, Ca ollia pe -
spicilla a, A ibeus jamaicensis, Dasyp oc a punc a a, and,
possibly, S l ilagus b asiliensis. All a e gene alis s wi hin
hei pa icula eeding niches and a e ypically o es -edge
species. Fo example, al hough Glossophaga so icina is a
nec a i o ous ba , i uses a b oade ange o plan species han
do o he nec a - eeding ba s.
Agou is and squi els a e conside ably mo e abundan a
La Sel a han in la ge ac s o lowland ain o es in he Ama-
zon Basin (pe s. obse a ion). Agou is and, p obably, o he
small o medium-sized oden s may ha e also bene i ed om
he sys ema ic killing o p eda o s by humans. An in e es ing
discussion o p eda o s in opical ecosys ems, wi h special
e e ence o La Sel a, is p esen ed by G eene (1988). He iden-
i ies one hund ed species o e eb a es as p eda o s a La
Sel a and independen ly concu s in a ibu ing he high popu-
la ion densi ies o many oden s cu en ly obse ed a La
Sel a and BCI o emo al o educ ion o p eda o popula ions
by humans.
The h ee common species o p ima es a La Sel a, A eles
geo oyi, Aloua a pallia a, and Cebus capucinus, all appea
o be mo e abundan now han hey we e in he 1960s and
ea ly 1970s. P ima e popula ions h oughou much o he Ca-
ibbean lowlands o Cen al Ame ica we e decima ed by an
epidemic o mosqui o-bo ne yellow e e du ing he ea ly
1950s (see Fishkind and Sussman 1987, and e e ences
he ein). Al hough da a a e no a ailable o he La Sel a e-
gion, one assumes ha his epidemic educed p ima e popula-
ions in he ese e. Fa ewe p ima es we e obse ed a La
Sel a be ween he la e 1960s and ea ly 1980s han in simila
ac s o p ima y o es elsewhe e in he Neo opics (pe s. ob-
se a ion). Al hough Mil on (1982) sugges s ha he howle
monkey popula ion on BCI had ebounded om he yellow
e e epidemic by 1970, ha eco e y was appa en ly no as
apid in he Sa apiqu egion, pe haps because o o he ac-
o s, including hun ing. P ima e popula ions a La Sel a, how-
e e , ha e ce ainly ebounded du ing he la e 1980s. Capu-
chin, howle , and spide monkeys can now be seen almos
daily and o en in la ge g oups. K. S one (pe s. comm.) es i-
ma ed in 1990 ha se en o wel e g oups o howle monkeys,
h ee o six g oups o spide monkeys, and ou o se en
g oups o whi e- aced capuchins we e on he g ea e La Sel a
p ope y. Howle monkeys a e he mos abundan . These e-
sul s ag ee wi h a p elimina y 10-day su ey conduc ed a La
Sel a in 1986 by Fishkind and Sussman (1987) al hough S on-
e 's numbe s o 1990 a e highe .
This inc ease in p ima e abundance may also be owed, in
pa , o a dec ease in he abundance o na u al p eda o s, espe-
cially ha py eagles
Ha pia
ha pyja). Cebid monkeys o he
gene a Aloua a, A eles, and Cebus cons i u ed mo e han one-
hi d o he p ey consumed by a pai o ha py eagles in Guy-
ana; wo- and h ee- oed slo hs, opossums, and agou is also
we e impo an p ey i ems (lzo 1985). Ha py eagles we e a e
in he Pue o Viejo egion as ea ly as he la e 1950s (Slud
1960) and a e now absen .
P ima es, as well as many o he species, undoub edly ben-
e i om comple e p o ec ion om hun ing a La Sel a. We
a e, pe haps, wi nessing a pe iod o luc ua ion in numbe s
o many p eda o y and p ey species because o he changing
deg ees o hun ing p essu e by humans. Because human popu-
la ions inc eased in he Pue o Viejo egion in he 1940s and
1950s be o e he es ablishmen o La Sel a as a ese e, one
would assume ha hun ing p essu e inc eased d ama ically.
Now ha La Sel a is well p o ec ed, hun ing p essu e has
been e ec i ely elimina ed om he ese e al hough poach-
ing con inues in B aulio Ca illo as a La Sel a in o he ea ly
1980s. I will be in e es ing o obse e he popula ion e-
sponses o p eda o s and he la ge p ey species. Complica ing
he abili y o de ec such changes is he ac ha indi iduals
o many species become less wa y when no hun ed and, hus,
a e mo e easily obse ed, which gi es he alse imp ession
o highe abundance e en hough numbe s may no ha e
changed.
His o ically, amilies in he Sa apiqui egion elied hea ily
upon local wildli e as a sou ce o p o ein. Tapi s, whi e-lipped
and colla ed pecca ies, and pacas a e highly p ized mea s and
a e among he la ges mammals o he egion. Medium-sized
mammals such as agou is, monkeys, and squi els (Sciu us
spp.) we e hun ed o a lesse ex en . Tapi s and pacas a e now
uncommon a La Sel a and in he gene al icini y e en whe e
adequa e habi a emains. Whi e-lipped pecca ies a e ex i -
pa ed om La Sel a; no sigh ings ha e been con i med on he
p ope y o se e al yea s al hough Slud (1960) ound hem o
be common he e in he la e 1950s. In he pas ew yea s, how-
e e , he popula ion o colla ed pecca ies has inc eased d a-
ma ically.
The ecen ex i pa ion o whi e-lipped pecca ies om La
Sel a may be a ec ing he na u e o he o es he e. Pecca ies
a e bo h majo seed dispe se s and seed p eda o s, and he
ac i i ies o la ge he ds could g ea ly a ec he o es plan s.
He d sizes o i y o mo e indi iduals a e commonly epo ed
o whi e-lipped pecca ies whe e hey a e no hea ily hun ed,
and hey end o concen a e hei ac i i ies in a eas o a o i e
ood esou ces.
La ge- and medium-sized mammalian p eda o s, including
jagua s, ocelo s, ay as, and coa is, ha e been pe secu ed by
humans in he Pue o Viejo egion as hey ha e been h ough-
ou he human-inhabi ed opical lowlands. These animals
we e killed o p o ec li es ock and c ops as well as o hei
pel s. O he p eda o s whose popula ions ha e been educed
in he Pue o Viejo egion, as well as h oughou hei anges,
include bushmas e s (Lachesis mu al and hawks and eagles
ha p ey on medium-sized oden s and p ima es. I is likely
ha he o he h ee species o ca s and he one i e o e ha e
also been a ec ed by hun ing and habi a des uc ion. Since
Cos a Rica's 1975 a i ica ion o he Con en ion on In e na-
ional T ade in Endange ed Species (CITES), he Cos a Rican
THE MAMMAL FAUNA 235
go e nmen has e ec i ely con olled illegal ade in wildli e.
P e iously, ca skins we e openly sold in ma ke s. Ca popula-
ions con inue o be se iously h ea ened by bo h habi a de-
s uc ion and hun ing. The o he species o ca ni o es a e, pe -
haps, nea ly as abundan now as in he pas whe e sui able
habi a exis s, bu sys ema ic s udies a e lacking.
Gian an ea e s p obably a e ex i pa ed in his egion and
h oughou mos o he coun y. O iginally, gian an ea e s
we e ound om Belize and Gua emala h oughou he low-
lands o Cen al Ame ica and opical Sou h Ame ica. Li le
is known abou hei o iginal dis ibu ion and abundance in
Cos a Rica. In he la e 1870s Als on (1879-82) epo ed ha
gian an ea e s we e a e in Cos a Rica and con ined o he
low,ho o es lands nea he coas . The e ha e been no epo s
o My mecophaga om his egion in ecen yea s, and one
p esumes ha hey a e now only ound in he mos emo e
egions o he coun y (Timm e al. 1989).
La Sel a, e en wi h i s connec ion, Pa que Nacional B au-
lio Ca illo, may be only minimally la ge enough o suppo
popula ions o p eda o y ba s o he sub amily Phyllos omi-
nae. The eigh een species o he phyllos omines ha I epo
he e o La Sel a p obably ep esen all he species expec ed
o occu he e. Some o hese la ge p eda o y species a e en-
coun e ed in much highe densi ies elsewhe e, in la ge ac s
o p is ine o es han cu en ly exis a La Sel a (pe s. obse -
a ion). La ge popula ions o p edaceous ba s we e epo ed
om La Sel a by LaVal and Fi ch (1977). Al hough all o
hese species a e s ill p esen in he ese e, mos a e no now
encoun e ed as equen ly as in he pas (pe s. obse a ion).
Appa en ly, his decline is he esul o he des uc ion o
much o he o es ha su ounded La Sel a and p o ided
mo e ex ensi e habi a o hese ba s. As wi h he la ge p eda-
o y ca s, hese ba s may be good "indica o " species in ha
hey may be among he i s componen s o disappea as a e-
sul o human dis u bance ( ig. 18.3). Thus, he connec ion o
La Sel a o he ecen ly expanded Pa que Nacional B aulio
Ca illo and he con inued e o o expand B aulio Ca illo
a e i ally impo an o he main enance o popula ions o
hese highly agile, p eda o y ba s, as well as o he mammals.
Human impac in his egion undoub edly has al e ed he
abundance o many species o mammals; some ha e inc eased
in abundance, whe eas many ha e dec eased in abundance.
Wi h he excep ion o gian an ea e s and whi e-lipped pecca -
ies, all species o mammals p esen a La Sel a be o e he
a i al and coloniza ion o he egion by Eu opeans ha e su -
i ed he e. La Sel a and he adjoining Pa que Nacional B au-
lio Ca illo a e la ge enough o p o ide sui able habi a o
mos species o mammals bu p obably no o la ge , highly
agile species such as gian an ea e s and whi e-lipped pecca-
ies. E e y e o should be made o p ese e as much o he
o es su ounding La Sel a and B aulio Ca illo as possible.
SUGGESTIONS FOR FUTURE RESEARCH
A e iew o he li e a u e on mammals a he La Sel a Biolog-
ical S a ion includes mo e han o y- i e o iginal published
con ibu ions. Mos deal wi h ecology and na u al his o y and
in e ac ions be ween mammals and o he bio ic componen s
o he en i onmen . Al hough mo e has been published abou
mammals a La Sel a han a o he Neo opical si es, excep
236 ROBERT M. TIMM
Fig. 18.3. The ca ni o ous ba Vampy um spec um has a la ge body and
is ound only in low numbe s. They a e monogamous and ha e only a
single young pe yea wi h ex ended pa en al ca e. Like he e es ial
p eda o s, i is among he i s mammal species o disappea wi h ag-
men a ion o ain o es s.
BCI much emains o be lea ned abou his complex auna and
i s componen species, When e iewing hese esea ch e-
po s. i is i onic ha one mus conclude ha mo e ques ions
ha e been posed han answe ed, In e es ingly. much mo e is
known abou he la ge mammals on BCI and e en a Manu
in Pe u han a La Sel a,
Nonmammalogis s migh wonde why ba s ha e been s ud-
ied so much a La Sel a (and elsewhe e in he Neo opics) and
oden s o o he g oups ha e been s udied so li le, in con as
o he oluminous li e a u e on empe a e oden s, Pe haps,
pa o he answe lies in he ela i e abundance o ba s in he
opics and hei ela i e ease o cap u e wi h mis ne s. Mos
mammal species a e noc u nal, wa y o humans, and ha e ex-
cellen hea ing and ision. Many o hem (especially ma supi-
als, p ima es, eden a es, oden s, and ca ni o es) a e pa ially
o wholly a bo eal in he opics, adding o he di icul y o
cap u e and s udy. The en i onmen a La Sel a has a h ee-
dimensional complexi y much g ea e han ha o empe a e
si es o e en o he opical si es, These di icul ies, along wi h
equen hea y ain all, ende di ec obse a ion, adio elem-
e y. and mos o he s anda d ma k- ecap u e echniques em-
ployed by empe a e mammalogis s di icul . Al hough mos
o he mammal esea ch a La Sel a has cen e ed on ba s. a -
emp s ha e been mace o sway o ne mammals. Some o
hese we e unsuccess ul. pe haps o some o he easons
men ioned abo e.
A spec acula opical o es such as ha ound a La Sel a
p esen s unlimi ed oppo uni ies o u u e s udy. This wo k
needs o be unde aken a se e al di e en le els, as ou lined
nex .
Faunal Su eys Coupled wi h Sys ema ic S udies o he
Mammal Species
Species o mammals undoub edly occu a La Sel a ha ha e
no ye been eco ded. These p e iously unde ec ed species
p obably include mo e han one species o ba and on•..o
mo e species o long- ailed oden s. This lack o he mos ba-
sic knowledge o wha species a e p esen is also ue o all
o he Neo opical si es. Resea che s qui e li e ally do no ha e
a comple e lis ing o he mammal auna o any New Wo ld
opical si e. In addi ion, knowledge o many o he small
mammals is so udimen a y ha undoub edly mo e han one
good biological species may be included unde a single name,
This con usion is almos ce ainly ue o O yzomys (Oligo -
yzomys) ul escens. Addi ionally, e en hough Cos a Rica is
one o he mos in ensi ely s udied coun ies in he Neo op-
ics, species o mammals new o science con inue o be disco -
e ed he e. Unde s anding o opical o es ecosys ems and
he abili y o make meaning ul compa isons be ween La Sel a
and o he si es a e hampe ed by incomple e knowledge o
he auna.
Habi a Requi emen s, Li e His o y S a egies, and
Rep oduc i e Modes
Essen ially no hing is known abou he habi a (and o he ) e-
qui emen s o mos species o mammals in he Neo opics,
especially he small and medium-sized species ha make up
mos o he auna. Gi en he now widesp ead a en ion o he
biodi e si y c isis in he opics. i is c i ical ha biologis s
ha e a be e unde s anding o he ecology o opical mam-
mals, De ailed knowledge o his so will be c i ical o he
p ope managemen o species and wildli e ese es in he u-
u e. One o he mos basic ques ions is How la ge an a ea
does a gi en species need o main ain a iable popula ion?
The answe has di ec implica ions o he size and shapes o
ese es,
Popula ion Biology and Communi y Ecology o
Neo opical Mammals
Knowledge o he s uc u e and dynamics o popula ions will
be c i ical o unde s anding how communi ies a e o ganized
and how opical sys ems di e om empe a e sys ems and
o unde s and all aspec s o e olu iona y biology heo y, S ud-
ies on a es o ep oduc ion, li e sizes. su i al a es. and
longe i y a e all easible and will p o ide conside able
insigh s.
Resea ch on mammals in he opics is en e ing an exci ing
phase. In es iga o s a e now pe ec ing he echniques needed
o wo k wi h hese animals. La Sel a's di e se auna p o ides
biologis s wi h nume ous oppo uni ies o explo e ecological
and e olu iona y ques ions. The ield acili ies a La Sel a a e
supe b. encou aging in e ac ions among scien is s and p o id-
ing excellen access o a ich auna.
ACKNOWLEDGMENTS
I hank he O ganiza ion o T opical S udies, especially R.
Bu e ield, D. and D. Cla k, L. A. McDade, C.
Eo
Schnell,
and D. E. S one o making my isi s o La Sel a o e he
yea s possible, pleasan , and p oduc i e. R. K. LaVal and
D. E. Wilson assis ed in de eloping he abundance indexes
THE MAMMAL FAUNA 237
p o ided in appendix 8, and I am g a e ul o hei inpu . S. E.
Ab ams, S. D. Ande son, A. P. B ooke, B. L. Clauson, and
K. E. S one p o ided cons uc i e e iew and insigh ul com-
men s on ea lie d a s o he manusc ip . Special hanks a e
due my wi e, Ba ba a Clauson, who has ably assis ed me wi h
my esea ch o e he yea s, and o p o iding bo h o he pho-
og aphs used he ein.
