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Group Title: Small mammals of the Cajas Plateau, southern Ecuador (FLMNH Bulletin v.42, no.4)
Title: Small mammals of the Cajas Plateau, southern Ecuador
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00099054/00001
 Material Information
Title: Small mammals of the Cajas Plateau, southern Ecuador ecology and natural history
Physical Description: p. 161-217 : ill. ; 23 cm.
Language: English
Creator: Barnett, Adrian
Florida Museum of Natural History
Donor: unknown ( endowment )
Publisher: Florida Museum of Natural History, University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1999
Copyright Date: 1999
Subject: Mammals -- Ecuador   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Spatial Coverage: Ecuador
Bibliography: Includes bibliographical references (p. 204-214).
General Note: Cover title.
General Note: Bulletin of the Florida Museum of Natural History, volume 42, number 4, pp. 161-217
Statement of Responsibility: Adrian A. Barnett.
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Bibliographic ID: UF00099054
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 43442549
issn - 0071-6154 ;


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Full Text

,z LET

of the



Adrian A. Barnett

Volume 42 No. 4, pp. 161-217



at irregular intervals. Volumes contain about 300 pages and are not necessarily completed in any one
calendar year.


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ISSN: 0071-6154


Publication Date: September 16, 1999

Price: $ 6.50


Adrian A. Barnett1


We captured 483 small mammals (19 species: 1 insectivore, 2 marsupials, 16 rodents) from sites
between 2700 and 4000 m on the Cajas Plateau, Azuay Province, Ecuador. Fifteen species were taken in
montane forest and 10 on the pAramo. Five species occurred in both habitats. One species was new to
science, three were recorded for only the second time and three records were major range extensions,
including one species new to Ecuador. Comparisons with other Andean sites show the species diversity of
small mammals on the Cajas Plateau to be among the highest recorded in the Andes. Reasons for this are
discussed. Data are presented on habitat preferences, reproduction, diet and pelage variation for each species.
Litter sizes at the small end of a species' range were consistently observed. Adversity selection (sensu
Southwood, 1977) at high altitude is suggested as an explanation. The natural history of several species is
reported for the first time since their original discovery.


Se capturaron 483 pequefios mamiferos pertenecientes a 19 species (1 insectivoro, 2 marsupiales y
16 roedores) desde sitios ubicados entire 2700 y 4000 m de altura en la Planicie de Cajas, Provincia de
Azuay, Ecuador. Se capturaron 15 species en la selva de montaiia y 10 en el paramo. Cinco species se
encontaron en ambos habitats. Una especie fue descrita como nueva para la ciencia, tres fueron registradas
s6lo por segunda vez y tres significaron extensions de rango mayor, incluyendo una especie nueva para el
Ecuador. La diversidad de species de pequefios mamiferos de la Planicie de Cajas es una de las mis altas
registrada para los Andes, de acuerdo a una comparaci6n con otras areas. Se discuten las razones de este
hallazgo. Se present informaci6n sore preferencias de habitat, reproducci6n, dieta y variaci6n de pelaje de
cada una de las species. El tamahio de camada observado se situ6 consistentemente en el rango inferior de
cada especie. Se sugiere como explicaci6n a esto filtimo la selecci6n por adversidad (sensu Southwood,
1977) a latitudes altas. La historic natural de varias de las species reportadas se describe por primer vez
desde su descubrimiento original.

I School of Life Sciences, Roehampton Institute. West Hill, London SW15 5SN, England, UK. and Akodon Ecological Consulting, 114
Petrie Avenue, Bryn Mawr, PA, 19010; USA.

Barnett, A. A. 1999. Small mammals of the Cajas Plateau, southern Ecuador: Ecology and natural history.
Bull. Florida Mus. Nat. Hist. 42(4): 161-217.



Introdu action ................................................................................................................................................ 16 2
A know ledgem ents .................................................................................................................................... 163
Site D escriptions........................................................................................................................................ 163
M methods ................... ............................................. .......................................................... ..... ....... 169
R results ................................................................ ....................................... ................... .............. 173
D iscussion........................................................................ .............................................. ............. 176
Species Ecology................................................................. ............................. ..... ................... 81
L literature C ited ............................................................................................................................. 20 4
Appendix 1 ....... ............................................................................ .. ................... 215
A pp en dix 2 ................................................................................................................................................. 2 16


Ecuador is one of the world's most biodiverse countries (Gentry, 1977;
Mitternneier et al., 1997), with approximately 340 species of mammals (Barnett
and Shapley, in prep. a)2. Despite this diversity, studies of Ecuador's mammals are
few compared to other Neotropical countries. Recent work on the small mammals
of Ecuador3 has made substantial contributions and yet mammals under 1 kg still
remain under-studied in Ecuador in comparison to neighboring countries. This is
illustrated by data from the Zoological Record. From, 1981 to, 1996, while this
organ gave the number of papers published on free-living native small mammals in
Colombia as 35, 50 for Peru, and 77 for Venezuela, Ecuador was limited to 20
This paper is a summary of work conducted on the small mammals of Cajas
region of Azuay Province, Ecuador. It was part of an Anglo-Ecuadorian research
initiative that worked first in the piramo (1981-84) and then in the cloud forest
vegetation of the region (1987-95) (see Barnett, 1986, 1988a; Barnett and Gretton,
1987). Mammal survey work was conducted between 1981 and 1987. For several of
the species covered here these reports are the first to appear on their natural history
since the species' original description.
Following these research periods, papers have been published on the bats
(Robinson, 1989), birds (Barnett, 1988b; King, 1989), invertebrates (e.g., Read,
1986), and plants (Aguilar and Espinoza, 1990-1991; Flores, 1993; Ramsay, 1992;
Ramsay and Oxley, 1996) of the Cajas region, and a number of papers on
individual species of non-volant mammals have appeared (Barnett, 1985, 1987,
1991, 1992, 1997a, b; Jenkins and Barnett, 1997; Barnett and Muleon, 1999).
These publications have not included all the non-volant small mammal species
known to occur on the Plateau. This paper is an attempt to rectify this and to

2This is substantially more than the 271 reported by Groombridge (1992). A copy of the species list may be
obtained by writing to the author.
3 (e.g., Figueroa and Albuja, 1983; Voss, 1983; Patton, 1984; Orcds, 1986; Voss, 1988; Carleton and
Musser, 1989; Albuja, 1991; Barnett, 1991; Best, 1991; Barnett, 1992; Parker and Carr, 1992; Voss, 1992;
Ragout and Albuja, 1994; Suarez et al., 1995; Albuja and Patterson, 1996; Bamrnett, 1997a; Jenkins and
Barnett, 1997; Schulenberg and Awbrey, 1997; Barnett and Muleon, 1999; Tirira, 1999).


present, in one place, comparative information on the habits and habitats of all the
small mammal species known to occur on the Cajas Plateau. A fuller analysis of
the biogeographic significance of Cajas will appear elsewhere (Barnett, in prep.).
Unless otherwise stated, taxonomy follows Wilson and Reeder (1993).


Luis Albuja and Gustavo Orc6s (Escuela Polytecnica Nacional, Quito), Sergio Figeroa (MAG,
Quito), Yolanda Kukabadse (Fundaci6n Natura, Quito), Tjitte de Vries (U. Cat6lica, Quito), Francisco
Escandon, Pablo Vientimilla and their staff (MAG, Cuenca), Agustin Rengel Barrera and the staff of
ETAPA (Cuenca), Noelia Montesinos and Comelio Montesinos (Tierra Viva, Cuenca), and the staff of the
Museo Ecuatoriana de Ciencias Naturales (Quito) for institutional and logistic support, kindness, advice, and
encouragement during fieldwork. Ivan Crouch, Philip Donlon, Carol Gordon, Trevor Iszatt, Fif Robinson,
John Shelvey, and Jon Wright for assistance in the field. Martin Perry and the current staff of the Mammal
Section, Natural History Museum, London (NHM), for loan of equipment, advice on specimens
identification, access to the collection and other technical help. I also thank the librarians of the Zoological
Society of London; the NHM; the Academy of Natural Sciences, Philadelphia; and the Smithsonian
Institution, Washington; as well as Guy Musser, Victor Pacheco, and Robert Voss, American Museum of
Natural History, New York; Michael Carleton and Craig Ludwig, Smithsonian Institution, Washington;
Bruce Patterson, Field Museum of Natural History, Chicago; and Jim Patton, University of California,
Berkeley. I should especially like to acknowledge Becca Shapley and Madeleine Prangley for their
forbearance and help with the manuscript.
This is a contribution from the Rio Mazin Project (UK) whose fieldwork has been funded by the
Birdlife International, British Ecological Society, Fauna and Flora International, Overseas Development
Administration, Percy Sladen Memorial Fund, Royal Geographical Society, Twenty Seven Foundation,
funds from the Natural History Museum, London, and many other sources. Publication of this paper was
funded by the Katharine Ordway Chair, Florida Museum of Natural History, University of Florida.

The Cajas Plateau, a formerly glaciated 25 x 27 km isolate of the Sierra de
Machangara of the western Cordillera in the Ecuadorian Andes, lies some 25 km
northwest from the town of Cuenca (2'52"S, 78'54"W) (see also Map 1). It is
represented on 1:50,000 maps CT-NV-3785-I, II, III and IV and CT-NVI-BI-3784-
IV, Instituto Geographico Militar, Quito. The plateau has a mean altitude of 3700
m, with peaks to 4138 m (Mt. Soldados). Colinveaux et al. (1997) give a detailed
vegetational history of the Cajas area for the last 13,000 years, based on
palynological analysis of cores from Lake Llaviuco. Our surveys included cloud
forest4 and pAramo vegetation types.
On the Cajas Plateau the treeline is at z 3400 m. Below this there are several
forest types. Areas of primary cloud forest are found in the Rio MazAn valley (see
Fig. 1). These were dominated by Ocotea aff. nectandra (Lauraceae), Podocarpus
sp. (Podocarpaceae), Clusia spp. (Clusiaceae), and Weinmannia fagaroides
(Cunoniaceae), interspersed with small areas of secondary forest dominated by

4sensu Patterson et al. (1998).







Recreacl6n Naclonal, Cajas


S National Boundary
'*"* Boundary of Azuay Province

Map 1: Location of Area Recreation Nacional Cajas in Ecuador.

-- KEY
Village/Town illi
Park Boundary Lake in
oRoad Paramo

Map 2: Sketch map of Area Recreation Nacional Cajas, showing study sites.

2 cm for 5Km


Figure 1. Primary cloud forest of the Rio Mazan valley (altitude 500-2800 m), showing one of the valleys
many waterfalls. Photograph, Adrian Barnett.


species of Miconia (Melastomataceae), including M. crocea. In the Zorracucho
Valley the slopes were covered with Myrtus-dominated secondary forest
interspersed by breaks of bamboo (mostly Chusquea spp.) and planted stands of
Pinus and Eucalyptus and an open valley floor covered with prominent clumps of
the graze-resistant shrub Barnadesia aff. arborea (Asteraceae). Both the MazAn
and Zorracucho valleys are on the eastern-facing side of the plateau. A third site,
Yakatuviana, was on the western-facing slope and was much moister than the
other two. Lamentably, the study site was highly disturbed, the formerly extensive
Clusia-dominated forest having been largely replaced with breaks of Chusquea
bamboo. Detailed botanical descriptions of specific localities are provided by
Fleming (1986) and Jacques et al. (1986), while Grubb et al. (1977), Patzelt
(1986), and Sanniento (1987) provided general and photographically illustrated
descriptions of Ecuadorian cloud forest structure and composition. Floristically, the
forests on the western side of the Cajas Plateau are similar to those at Pasachoa
described by Jorgensen and Valencia (1988) and those to the south described by
Madsen and Ollgaard (1994). Because they face the Pacific Ocean, the forests on
the western slope of the Andes (e.g., at Yakatuviana) are much moister and
wanner than their eastern counterparts (see Grubb, 1977; Patzelt, 1986; Sarnniento,
Between the forest and the paramo, at 3500 in, is a shrubby ecotone some
100 m wide. Above this is pAramo moorland (see Fig. 2), corresponding to the
'grassy piramo' division of Cuatrecasas (1968) and Harling (1979). Within this
there are three broad subdivisions; pajonal (grass paralno) from 3400 m to 4000 m
and, above this, shrub and cushion pAramo (=yareta)'. The third division, quenoa
forest, occurs in sheltered locations in both piramo belts (see Fig. 2). Pajonal and
yareta are both mosaics of vegetation types, whose local composition is influenced
by drainage and exposure (Smith, 1977; Van Der Hammen and Cleef, 1986).
In pajonal several different micro-environments are found. At any altitude the
most exposed areas are dominated by plants with low-growing and cushion-like
growth forms (see Acosta-Solis, 1966; Van Cleef, 1981). In sheltered areas there
are quenoa forests dominated by the short, twisted trunks of Polylepis (Roseaceae)
trees (see Fig. 3). The edges of the forests frequently have dense stands of
composite shrubs (Gynoxis and Diphlostephium) that act as windbreaks. These
near windless, moist environments have a dense cover of ground-living and
epiphytic bryophytes. Along with gullies ('quebradas'), rock outcrops and
riverbanks, these forests fonn mesic islands in an otherwise wind-dried
environment (see Simpson, 1979a; Barnett, 1992; Ramsay, 1992; Ashe and
Leschen, 1995; FjeldsA and Kessler, 1996). In the quebradas there is less moss
cover and characteristic vegetation includes Bomarea vines (Alstroemeriaceae) and
shrubs of the genera Cavendischia (Ericaceae), Baccharis (Asteraceae) and

5 though this is a common name of a paramo plant (Azorella) in the altiplano of Peru, Ramsay (1992) has
used it to describe the community of cushion plants of which Azorella is characteristic, a use that is followed

:Y~ ~

Figure 2. Paramo showing pajonal grassland and quenoa forests. Note the small extent of the forests and their restriction to sheltered
areas. Near Lake Torreadora (4000 m). Photograph, Adrian Barnett.



Figure 3. Interior of a quenoa forest, showing tangled nature of the Polylepis trees, the low canopy and the
thick mossy groundcover (near Lake Torreadora, 4000 m). Photograph, Adrian Barnett.


Calceolaria (Scrophulariaceae). Rock outcrops (see Fig. 4) frequently shelter
Calceolaria spp. and members of the ericaceous genera Maclenia, Gautheria and
The pAramo of Cajas has been quantitatively analyzed by Ramsay (1992),
who distinguished six vegetation associations. A botanical summary of those
relevant here are presented in Table 1.
Around 80% of the Cajas Plateau lies within the boundaries of the Area
Nacional de Recreacion, Cajas (ARN) (see Huber, 1979). As such it has received
some wardening and protection since the ARN's inception in 1976. Nevertheless,
some parts of the ARN are subject to intense human pressure, including cattle
grazing and deliberate fire-setting as part of local range-management practices
(Huber, 1979; Ramsay and Oxley, 1996; see also Knapp, 1991). Other areas are
popular with fishermen and few areas remain fully undisturbed (Barnett, 1997a, b).
As with pAramos elsewhere (Ralph, 1978; Smith, 1972; Baruch, 1979; Smith,
1981; Van Der Hammen and Cleef, 1986), exposure and wind chill can be
important physical constraints on the fauna and flora of Cajas. There is no
permanent snowline, though snow may fall on the higher peaks at any time of year.
Weather is frequently cold and overcast and winds often gust to over 60 kmph (see
Ramsay, 1992; Barnett, pers. obs.). Daily temperature ranges from -60C at night to
270C during the day. This exceeds the annual mean temperature variation. There
has been no long term collection of meteorological data on Cajas, but 30 years of
records for Cuenca (2000 m) show that rainfall is highest between March and May.
The dry season is from June to January in southern Ecuador (Ramsay, 1992), with
the lowest rainfall in Cuenca between June and September (Meteorological Office,


Prior to the fieldwork covered in this report, the small mammals of the Cajas
Plateau were effectively unstudied (see Huber, 1979). The main aim, therefore, was
to obtain the most representatively accurate sample of the species present over the
whole plateau, rather than conduct detailed quantitative investigations of particular
areas within the ARN. This influenced the operational regime deployed. Study sites
were chosen to cover the full range of habitat types in the ARN and adjacent areas.
Eleven sites were visited (see Map 2). Information on the altitude and habitat types
of these is given in Table 2. The rugged terrain made the use of regular trapping
grids ineffective. Accordingly, traps were placed at runs, holes, and other sites of
small mammal activity (see Twigg, 1975a; Gurnell and Flowerdew, 1990; Barnett
and Dutton, 1995). This provided less quantifiable results but probably increased
trapping success, an acceptable compromise in little-known areas (Barnett and
Dutton, 1995; Barnett, 1996; Stork and Davies, 1996). A mixture of traps were
used, including live (Longworths, Shermans, and Havaharts) and snap traps



Figure 4. A rock outcrop with attendant Calceolaria shrubs. Near Lake Luspa (3700 m). Photograph,
Adrian Barnett.


Table 1. Summary of the botanical characteristics of paramo-sub-types at Cajas (modified from Ramsay,

Name of paramo sub-type range Floral characteristics

Viola humboltii pajonal 3500-3600 Dominated by Calamagrostris bunchgrass with scattered
m shrubs and low-growing rosette plants including Viola
humboltii, Azorella pedunculata, Halenia weddelliana,
Geranium reptans, G. multipartium and G. sibbaldoides.

Paspalum pajonal 3500-3600 More open grassland than V humbolti-grassland with a higher
m proportion of Paspalum tuberosum (Graminae), mixed with
Lycopodium clavatum (Bryophyta: Lycopodiaceae), Eryngium
humble (Asteraceae), Hypochaeris sessiliflora (Asteraceae)
and Hypertcum (Guttiferaceae).

Shrubby piramo 3800 m Characterized by composite shrubs of the genus Baccharis (inc.
B. alpinum), Diphlostephium (inc. D. hartwegii) and
Chusquiragua insigms, and the herbs Gentianela diffusa
(Gentianaceae), Baccharis genistelloides and Castilleja
pumila (Scrophulariaceae).

Yareta Cushion pAramo 3800 m Dominated by plants with low-growing and cushion-like growth
forms, of which Hypochaeris sessiliflora and Werneria
nubigena (Asteraceae), Phyllactis rigida (Valerianaceae), the
gentians Gentiana sedifolia and Halenia mayeri and Is6etes
andina (Pteridophyta: Isoetaceae) are characteristic. Azorella
compact (Umbelliferae) is also present.

(Nippers, Little Nippers, Self-sets, and locally available metal snap-traps of
Chinese manufacture)6
Traps were flagged to aid re-location and staked and tied to avoid removal by
scavengers (see Gurnell and Flowerdew, 1990; Barnett and Dutton, 1995). In
especially dense and difficult terrain, where the visual horizon was less than the
distance between the traps (notably in quenoa forests and cloud forest), a white
cord was run between traps at eye height (termed 'festooning' by Barnett and
Dutton, 1995) to speed trap location and so minimize vegetation damage and

6 Manufacturers' information:
Havahart traps, Havahart Ltd., Box 551, Ossining, New York, USA;
Longworth Live traps, Penlon Ltd., Radley Road, Abingdon, Oxfordshire, OX14 3PH, UK;
Sherman traps, H.B. Sherman Inc., P.O. Box 20267, Tallahassee, FLA 32316, USA;
Nippers and Little Nippers, Procter Bros. Ltd., Pantglas Industial Estate, Bedwas, Newport, Gwent NP1
8XD, UK;
Self-sets, Falcon Works, Hanworth Road, Sunbury-on-Thames, Surrey, TW16 5DE, UK.


Table 2. Site descriptions.

Locality Altitude Description

Lake Torreadora 4000 m Area of Viola pajonal with frequent quenoa forests and some areas
of marshland Paspalum pajonal and yareta. Forests and grassland
subject to heavy visitor pressure. Frequent waterfalls and streams.

Athiovacu 3900 m Small, steep-sided protected valley filled with shrubby pAramo and
quenoa forest. Little disturbance.

Lake Quinoascocha 3800 m Very exposed locality. Viola pajonal, quenoa forest, Gynoxis
clumps and scrub-dotted scree. Little disturbance.

Chapiurcu 3750 m Wide shallow valley filled with remnants of once-considerable
quenoa forest, now greatly fragmented by hearing activities and

Lake Luspa 3700 m Viola pajonal, yareta, marsh and quenoa forest. Disturbance by
grazing, recreational fishing, and tourism.

Ventanas 3650 m Heavily grazed grassland, formerly Viola and Yareta, but with
close-cropped grass tussocks all most rosette plants trampled. Some
quenoa forest and rock outcrops.

Chirimanchi 3450 m Stunted trees and shrubs at piramo/montane forest ecotone.

Lake Llaviuco 3100 m. Secondary montane forest on valley sides. Grazed valley floor.
(Zorracucho valley) (eastern Reedy marsh around lake.
side of

Rio MazAn valley 2700 m Primary Podocarpus and Ocotea-dominated montane forest, with
(eastern some secondary forest and areas of grazed grassland.
side of

Yakatuviana 2400 m Bamboo (Chusquea spp.)-dominated montane forest. Some small-
(western scale agriculture.
side of


associated reduction of trap success. Bait was a mixture of porridge oats, cooking
oil, banana and water. Sardines, tuna, trout flesh, sausage, or meat were used for
faunivorous species. Snap-traps were checked twice a day from 0700 to 0800 hrs
and from 1200 to 1300 hrs. Live-traps were checked from 0600 to 0700 hrs and
from 1700 to 1800 hrs. Data collection methods and specimen preparation
followed Corbet (1968). Methodology for capture-mark-release studies followed
Twigg (1975b). Field work took place between June and October in 1981; 1983,
1984, 1986, and 1987.
All specimens were initially identified by Martin C. Perry of the Mammal
Section, Natural History Museum, London. Specimens of Microryzomys were
subsequently re-identified by Michael Carleton and Guy Musser (see Carleton and
Musser, 1989), and some specimens of Oryzomys were subsequently re-identified
by Guy Musser (AMNH), while Rob Voss (AMNH) re-identified some specimens
of Thomasomys. Synonymies for all species collected are given in Appendix 1.
Unless there is a more recent taxonomic treatment, these are based on Wilson and
Reeder (1993).


General trapping.-The total trapping effort was 5942 trap nights. This
consisted of 4178 trap nights in paramo and 1764 trap nights in cloud, pine, and
Eucalyptus forests (1302 at MazAn, 402 at Llaviuco, 60 at Yakatuviana). Of these
1666 trap nights were expended live-trapping for fishing mice (Ichthyomyini), 130
were spent live-trapping at Llaviuco and a further 240 spent studying movement
patterns of rodents inhabiting quenoa forest.
Four hundred and eighty three individual small mammals were captured. Of
these 468 were rodents (16 species, 8 genera), twelve were insectivores (1 species)
and three were marsupials (2 species, 1 genus). Of these 440 were made into
museum specimens. In addition, 53 examples of non-target species were also
caught; rufous ant-pitta Grallaria rufula (3), plumbaceous sierra finch Phrygilus
unicolor (1), bar-winged cinclodes Cinclodes fuscus (6), frogs of the genera
Atelopus, Eleutherodactylus, and Gastrotheca (17), and invertebrates (mainly
gastropods, phasmids, and coleoptera, 26). Unlike lowland sites (Barnett and da
Cunha, 1994), bait removal by ants at Cajas was minimal (less than 1% of trap-
nights), but 8% of trap-nights were lost to rain disturbance (traps set off or bait
washed off). One site was subject to disturbance by large mammals.
General snap-trapping.-In 3906 trap nights, 229 specimens of 18 species
were caught. The low overall trap success (5.86%) masks great inter-site and inter-
habitat variability in trapping success (see Table 3)7
Trapping for fishing mice.-Three specimens of one ichthyomyine species,
Chibchanomys orcesi, were captured in live traps. In addition, three shrews,

data is given for all sites except Athiovacu, for which trapping data has been lost.


Table 3. Snap-trap success and small mammal community composition between localities and habitats.

Locality, % Trap %Akodon
Habitats. success mollis Species captured

Lake Torreadora 68.6 77.3
Quenoa 11.1 54.6 C. montivaga (quenoa,
Viola pajonal 07.0 85.7 quebradas)
Paspalum pajonal 12.12 75.0 A. mollis (all, except bog)
Rock outcrops 12.7 62.5 M. altissimus (quenoa, rock
Quebradas (gullies, 10.5 62.7 outcrops)
streamsides) P. haggardi (rocks, paspalum)
Shrub patches T. gracilis (quenoa)
Quaking bog T. pyrrhonotus (quenoa)
House S. inopinatus (quaking bog)

Quenoa no information
Viola Pajonal
Lake Quinoascocha 07.1 73.9
Paspalum pajonal 09.33 100 C. montivaga (quenoa)
Shrub patches 06.3 80.0 A. mollis (all)
Quenoa 06.9 41.7 M. altissimus (quenoa,
paspalum pajonal)
T. cinnameus (quenoa, shrub
T. gracilis (quenoa)

Chapiurcu 12.7 73.33
Quenoa 12.4 100 C. montivaga (quenoa)
Shrubs 10.1 80.0 A. mollis (quenoa, shrubs)
Riverbank 20.0 40.0 M. altissimus (quenoa,
P. haggardi (rocks)
S. inopinatus (riverbank
T. gracilis (quenoa)

Lake Luspa 22.3 55.5
Quenoa 19.4 32.0 C. montivaga (quenoa,
Viola pajonal 26.0 50.0 quebradas)
Paspalum pajonal 08.7 50.0 A. mollis (all, except stream)
Rock outcrops 75.0 44.5 C. orcesi (stream)
Quebradas 19.0 38.5 M. altissimus (quenoa, rocks)
Shrub patches 14.8 100 P. andium (rocks)
T. gracilis (quenoa, rocks)
T. pyrrhonotus (quenoa)

Ventanas 13.75 47.6
Quenoa 12.5 33.33 A. mollis (pajonal, quenoa)
Viola Pajonal 15.0 100 M. altissimus (quenoa)


Chirimanchi 15 50
Streamside C. montivaga
quebrada A. mollis
M. altissimus

Lake Llaviuco 11.2 26.46
(Zorracucho valley) A. mollis (marsh, grassland,
Secondary forest 09.2 0 Eucalyptus/Pinus)
Marsh 02.5 33.0 A. orophilus (forest)
Grazed grassland 02.5 100 C. orcesi (streams)
Riverbank 05.0 0 M. altissimus (forest)
Eucalyptus grove 01.7 100 M. minutus (forest)
0. destructor (forest, hut,
T. aureus (forest, marsh)
T. baeops (forest)
T. gracilis (forest, marsh)
T paramorum
T pyrrhonotus (forest)
Rio Mazan valley 7.0 51.4
Podocarpus forest 5.6 0 C. caniventer (forest)
Riverside 6.9 56.1 C. tatei (forest)
Grazed grassland 9.6 75.0 C. montivaga (forest)
A. mollis (grassland)
A. orophilus (forest)
0. albigularis (forest)
T. aureus (forest)
T. baeops (forest)
T. paramorum (forest)

Yakatuviana 11.1 64.3
Primary forest 02.5 0 A. orophilus (secondary forest, field)
Secondary forest 09.7 28.6 0. albigularis (primary and
Agricultural field 12.5 100 secondary forest)
T. paramorum (secondary

Cryptotis montivaga, and 232 individuals of 6 rodent species (200 A. mollis, 4
Microryzomys altissimus, 1 Oligoryzomys destructor, 8 Oryzomys albigularis, 1
Sigmodon inopinatus, 8 Thomasomys gracilis) were caught in 1666 trap nights (see
Barnett, 1997a: table 1). Fourteen non-mammals were also captured. While total
mammal trap success was 14.28%, trapping success for ichthyomyines was very
low (0.21%).
Movement patterns of quenoa-forest inhabiting rodents.-Four lines of
twelve Longworth live traps were set for five nights, with equal proportions of
traps in the quenoa and adjacent grassland. Inter-trap distance was 10 m,
transects were = 100 m apart. Twelve individuals from four species (Akodon
mollis, Microryzomys altissimus, Thomasomys cinnameus, Thomasomys gracilis)
were caught a total of 36 times in 240 trap nights. The results of the capture-mark-


Table 4. Live-trapping data showing distances traveled and percentage occupancy of dense cover.

Number of Distance (m) between inter-trap % records in
Animals captured Recaptures Recaptures1'2 Distance3 (m) Dense cover4

A. mollis 1 2 138,0 138 100
A. mollis 2 4 40; 110;34,0 110 50
A. mollis 3 4 112, 0; 0; 41.6 112 25
A. mollis 4 0 0
M. altissimus 1 0 100
M. altissimus 2 3 0; 20; 31.2 31.2 33.3
T. cinnameus 1 2 0; 10 10 100
T. cinnameus 2 3 11.5, 0; 0 11.5 100
T. gracilis 1 1 152.3 152.3 100
T. gracilis 2 0 100
T. gracilis 3 1 10 10 100
T. gracilis 4 4 10; 0; 20; 11.7 20 100

1: 0 indicates caught in the same trap as proceeding record.
2: calculated by direct line measurement between traps.
3 includes all habitat types, value direct line measurement
4 calculated by number of records in each habitat type.

recapture study are given in Table 4 and discussed on a species-by-species basis
under the species headings.
Trapping in Pine Plantations and in Eucalyptus plantations and live-
trapping, Zorracucho valley.-Sixty nights snap-trapping in pine and Eucalyptus
plantations near Lake Llaviuco yielded a single immature male Akodon mollis. The
open, grazed floor of the valley was regularly disturbed by cows and children. Live-
trapping there with Longworths for 130 nights caught three A. mollis.


Trapping.-Capture rates in Cajas were much higher than those generally
expected of the species-rich, numbers poor tropical lowland communities, where
rates of 0.5-2.5% are common (Voss and Emmonsl996) Similar success rates to
those in Cajas have been reported in other Andean communities (e.g., Meserve et
al., 1982; Murfia and GonzAlez, 1982; Meserve et al., 1991). Such low trapping
rates as have been reported for Andean small mammal communities (e.g., Zunifiga
et al., 1983, 0.33% and 0.43%; Pdfaur and Diaz-de-Piscual, 1985, 3.75%; Lopez-
Arevalo et al., 1993, 4.31%; Durant and Diaz, 1995, approx. 2.68%) are most

8 though this pattern is not universal. Studies by James Patton in western Brazil obtained trap success
between 5 and 12% in terra fire forest, and even higher in virzea, though these numbers varied seasonally
(J. Patton, pers. comm., 1998: Patton et al., 1994).


probably methodological artifacts resulting from the use of brief sampling periods
(Zunifiga et al., 1983; Pefaur and Diaz-de-Pascual, 1985), the exclusive use of live-
traps (e.g., Lopez-Arevalo et al, 1993, which generally give a lower capture success
rate than snap traps [see Hansson and Hoffmeyer, 1973; Barnett and Dutton,
1995]), or both (Durant and Diaz, 1995). A number of authors (e.g., Emmons,
1984; Barnett and da Cunha, 1994; Voss and Emmons, 1996; Barnett and da
Cunha, 1998a) have reported positive correlations between trapping success and
soil fertility. Accordingly, the high trap success at Cajas (and elsewhere in the
moist Andes) may be attributable to the fertile volcanic-derived soil of much of the
mountain chain (see Kennerley, 1980; Owen, 1990).
Trap success was highest in quenoa forest and in rock falls. It is unclear if
this is because there were more animals there, or because the physical structure of
the habitat assisted entrapment (moss-covered floors easily reveal small mammal
trails-see Carey and Witt, 1991; Malcolm, 1991) and rock jumbles limit the
places for a small mammal to run (Barnett and Dutton, 1995). The low capture
rates in pine and Eucalyptus plantations are in line with data from similar
plantations in other countries (e.g., Friend, 1982; Happold and Happold, 1987;
Mitchell et al., 1995; Stanko et al., 1996; Zejda and Nesvadbova, 1996) and may
be due to the poorly developed groundcover and leaflitter layers and the dearth of
insect and seed forage available (see Allen et al., 1995; Ogden et al., 1997).
The low capture rates for ichthyomyines was not unexpected as trap-shyness
seems to be typical for the group. As examples, Anthony (1921: 2) reported that
having obtained the type of Ichthyomys tweedi, "although traps were set out in
every suitable locality ... no additional specimens were secured" (see also Anthony,
1929: 1); Andrea Pogson and Caroline Lees trapped specifically for ichthyomyines
and caught one in 700 trap nights (see Barnett, 1997a), and Ochoa and Soriano
(1991) reported very low success trapping for Neusticomys mussoi in Venezuela, as
did Durant and Diaz (1995) for Ichthyomys hydrobates.
Live trapping results from the quenoa forest and pAramo showed that, of the
12 animals caught, only four were trapped away from dense cover. A. mollis
showed the greatest mobility, with a mean maximum inter-trap distance (mn.mr.i.d.)
of 120 m over the study period (see Table 4). Thomasomys gracilis was also quite
mobile, with an m.m.i.d. of 45.5 m. T. cinnameus appeared to move little (m.m.i.d.
of 10.75 m). These singularities may reflect differences in diet and the dispersion
and patchiness of food resources, which has often been shown to influence the
ranging patterns of small mammals (e.g., Dalby, 1975; Delany, 1986; R.G.
Anthony et al., 1987; Happold and Happold, 1989, 1992; Canova and Fasola,
1993; Ellis et al., 1997). If this is occurring with Akodon and Thomasomys in the
quenoa forests of Cajas, it is doing so counter-intuitively, for Akodon is a generalist
(browse, fungi, seeds, and insects), while Thomasomys is predominantly
frugivorous (see Nowak, 1991; Lopez-Arevalo et al., 1993). This would lead one to
predict the reverse pattern of range sizes for the two genera. Larger sample sizes
and quantified records of prey-base abundance and distribution are required before
valid conclusions can be drawn.


General Ecological Structuring

It is instructive to compare the number of species present at Cajas with those
reported by other workers from other areas in the Andes. Comparisons are often
difficult because of methodological differences'- which can lead to differences in
the proportion of the catch each species represents (see Nichols, 1986).
Comparisons are therefore restricted simply to the number of species, rather than
any more refined standard of comparison.
The small mammal species diversity of the cloud forest zone of Cajas (2
marsupials, 1 insectivore, 12 rodents) compares favorably with similar high
altitude forest study sites in Venezuela (Diaz-de-Piscual, 1984, 1994; Pdfaur and
Diaz-de-PAscual, 1985; Durant and Diaz, 1995), Colombia (Durant and PMfaur,
1984; Lopez-Arevalo et al., 1993), Ecuador (Albuja and Luna, 1997), and Peru
(Dorst, 1958; Patton, 1986; Pacheco et al., 1993) and with the more xeric Peruvian
communities Pearson (1951) and Pearson and Ralph (1978) studied (see Table 5).
However, the diversity of small mammals in the pAramo of Cajas (1
insectivore, 9 rodents) is considerably higher than other comparable sites (Dorst,
1958; Durant and Pdfaur, 1984; Lopez-Arevalo et al., 1993; Durant and Diaz,
1995), and close to that reported by Pearson (1951) and Pearson and Ralph (1978)
(see Table 5). With the exception of Patton (1986) and Pacheco et al. (1993), the
combined small mammal species diversity for montane forest and pAramo is higher
in Cajas than other studied sites (see Table 5). This may be due to the special
biogeographical position of Cajas, which embraces the northern-most ingress of the
genus Phyllotis and the southern-most extensions of the genera Sigmodon and
Comparison with Cotgreave's and Stockley's (1994) compilation of species
diversity data for 25 small mammal communities around the world shows that
Cajas is highly diverse. Only two of their listed sites had numbers of species
comparable to Cajas'" However, Cajas' small mammal species diversity is between
105% and 51% of that collated by Voss and Emmons (1996) for 10 lowland
rainforest sites in the Amazon Basin. The numbers of rodent species are broadly
similar (range 11-27), and the differences are largely due to the absence of

Se.g., see Neal and Cock (1969), Schwan (1986) for trap type; Willan (1986) for bait; Hansson (1975) for
quadrat size; Vickery and Bider (1981) for weather; Gurnell and Langbein (1983) for trap positioning; Tew
(1987); Reynolds and Gorman (1994 ) for seasons and Shore and Yalden (1991), Tew et al. (1994 ) for
minute variations in technique
10 Formerly considered to occur no further south than Loja Province, Ecuador (see Barnett, 1992), Cryptotis
has now been recorded in northern Peru by Vivar et al. (1997). Sigmodon is much more speciose in the
northern part of its range (Voss, 1992). Cajas is the southern-most extention ofS. inopinatus (Voss, 1992),
with only Sigmodon peruanus from northern Peru occurring further south (Pacheco et al., 1995).
11 though the scales are probably different. Though area is not always given, none of the sites used either by
Cotgreve and Stockley or by Voss and Emmons would appear to cover an area comparable to Cajas' 25x27


Table 5. Comparative species diversity for Andean small mammal communities.

Number of species
Location I=insectivores,
Study and habitat type (situation, country, altitude) R=rodents)

Montane forest

Diaz-de-Pascual, 1984 north-eastern Venezuela, 500 m 0 M, 0 I, 8 R
Durant and Diaz, 1995 north-western Venezuela, 1890- 0 M, 1 I, 9 R
2600 m
Pefaur and Diaz-de-Pascual, 1985 north-eastern Venezuela, 1000- 16 and 17 species
2500 m
Durant and P6faur, 1984 Merida, Colombia, 1640-2500 m 1 M, 1 I, 5 R
Lopez-Arevalo et al., 1993 Cord. Oriental, Colombia, 2380- 2 M, 1 I, 5 R
2900 m
Zufiinga et al., 1983 Cord. Occidental, Colombia, 2000- 1 M, 0 I, 2 R
3000 m
this study Cajas, Ecuador, 2400-3100 m 2 m, 1 I, 12 R
Albuja and Luna, 1997 Cord. de C6ndor, Ecuador, 1000- 1 M, 0 I, 3 R
1600 m
Patton, 1986 southern Peru, 2000-3000 m 1 M, --, 10 R
Pacheco et al., 1993 central Peru, 1100-1880 m 3 M, --, 9 R

High-altitude grassland

Durant and Diaz, 1995 north-western Venezuela, 3000- 0 M, 1 1, 3 R
3950m 0M, 1 I1, 3R
Durant and P6faur, 1984 Merida, Colombia, 3300-3600 m
Lopez-Arevalo et al., 1993 Cord. Oriental, Colombia, 3000- 0 M, 1 I, 4 R
3380 m
this study Cajas, Ecuador, 3300-4000 m 0 M, 1 I, 10 R
Dorst, 1958 Andes, central Peru (altiplano), 0 M, 0 I, 8 R
4000 m
Patton, 1986 Andes, southern Peru (puna), 3500- -, --, 13 R
5000 m
Pearson, 1951 Altiplano, southern Peru, 3000- 5-14 R per site
5000 m
Pearson and Ralph, 1978 Pacific-facing slope, northern Peru 1 M, O I, 16 R
(puna), 3500-4500 m
Pacheco et al, 1993 Amazon-facing slope, central Peru, 1 M, -, 8 R
2800-3450 m

marsupials at higher altitudes compared to those in the Amazon (where there are
between 5 and 12 marsupial species per site)"2
In Cajas, small mammal species diversity was slightly higher in the cloud
forests than in the paramo (15 vs. 10 species), with 9 and 4 species, respectively
being unique to each zone. The greater species numbers in cloud forest are
probably a consequence of within-habitat variation attributable to the greater

12see also Pacheco et al. (1993).



variety of habitat types present at each locality (see Veillon, 1965; Pearson and
Ralph, 1978; Gibb, 1981). It should also be noted that there is considerable
between-site variation, with the three cloud forest areas sampled being very
different in their dominant plant communities; MazAn was almost exclusively
primary forest, Zorracucho was mostly secondary forest and Yakatuviana was
much moister than either due to its location (the Plateau's Pacific-facing slope).
This diversity may well have increased the pool of small mammal species available
for sampling.
Above 2400 m, there are well-marked differences in plant communities that
comprise the paramo grassland sub-types (see above and Table 1). However, the
greatest number of small mammals was caught in sheltered areas (284 of 301
animals). Only four species (Akodon mollis, Microryzomys altissimus, Phyllotis
andium and P. haggardi) were caught in grassland pAramo and only the latter two
were caught uniquely there. No animals were caught on yareta pAramo. Yareta is
highly exposed, lacks cover, and the surface, provided by the continuous cover of
rosette plants, is hard and unyielding. This may have made it unattractive to small
mammals. It would seem that the higher-altitude living rodents of Cajas favor
habitats provided by quenoa forests, vegetated quebradas, and streamside
vegetation. These probably provide both more shelter and more insect prey,
significant factors when the diet so many high-altitude rodents includes insects (see
Glanz, 1982; Lopez-Arevalo et al., 1993; Cotgreave and Stockley, 1994). Such
habitats are both micro-geographically restricted and compositionally quite
uniform (see Acosta-Solis, 1966; Simpson, 1976a; Ramsay, 1992), compared to
montane forest (Veillon, 1965; Grubb, 1977; Grubb et al., 1977). Thus, from the
small mammal perspective, the pAramo may be less variable than it is to the eye of
a botanist.
Higher species diversity at lower elevation is a common pattern for many taxa
(see Rahbek, 1995) and has been documented in the montane small mammal
faunas of Mexico (Fa et al., 1990), Peru (Pacheco et al., 1993), Malaysia (Medway,
1972), New Guinea (Laurie, 1952) and Africa (Misonne, 1963; Rowe-Rowe and
Meester, 1982). However, this phenomenon is not uniform; the opposite trend
having been observed in the western slope of the Peruvian Andes (Pearson and
Ralph, 1978) eastern slope of the Peruvian Andes (Patton, 1986), on the eastern
slope of the Venezuelan Andes (Pdfaur and Diaz-de-PAscual, 1985), in the
Ethiopian highlands (Yalden, 1988), and in Texas (Owen, 1990). Some studies
(e.g., Heaney et al., 1982, in the Philippines; Happold and Happold, 1992, in
Africa; Patterson et al., 1998, in Peru13) have found no difference in small mammal
species diversity between altitudes, while others (e.g., Yu, 1994) have found
greatest species diversity at intermediate altitudes. Intra-locality differences in

13 For a 3 km altitudinal transect from Amazon lowlands to the Puna grasslands of south-eastern Peru
Patterson et al. (1998), found that the number ofmurid species showed little relationship to altitude.
However, over the part of the altitudinal and habitat range comparable to the habitats present in Cajas, the
number of all species of small mammal (marsupials and all rodents) was greater in montane forest than in
high altitude grasslands (see data in Pacheco et al., 1993).


species diversity can have ecological and/or historical explanations (Endler, 1982;
Vuilleumier and Monasterio, 1986; Voss and Emmons, 1996). Explanations for the
current differences between sites could include the way habitat complexity is
distributed over the studied altitudinal range and the nature of annual climatic
differences, are probably the keys to this apparent lack of phenomonological
homogeneity (see Happold and Happold, 1992; Patterson et al., 1998). Historical
explanations could include the altitude, glacial history, and past inter-connections
with other areas (Vuilleumier and Simberloff, 1980; Patterson et al., 1998; J.
Patton, pers. comm.).
Tropical ecosystems are generally characterized by high species diversity,
high equatibility, low numbers of individuals, temperate ones by with lower species
diversity, skewed equatability, and higher numbers of individuals (Glanz, 1982;
O'Connell, 1982; Claridge et al., 1997). The relatively low mean trap success
(9.76%), higher equitability (the numerically dominant species, A. mollis,
comprising 29.8% of specimens), and higher species diversity of the montane
forest small mammal fauna contrast with that of the paramo (mean trap success
23.3%; 75.5% A. mollis). These indicators show that, while the montane small
mammal communities may be structured like tropical systems, that of the pAramo
more closely resembles a temperate situation (see O'Connell, 1982). This has also
been observed for certain groups of Andean plants (Simpson, 1974), reptiles and
amphibians (Duellman, 1979), and birds (Vuilleumier, 1986).
Reig (1986) compiled a list of rodents living in one or more of the 23 paramo
areas of the northern Andes recognized by Vuilleumier and Simberloff (1980). He
did not make comparisons between the localities nor provide separate lists for
various sites, but listed 27 species overall. With nine paramo species, Cajas would
appear to have a good representation of northern Andean high-altitude rodents.
The taxonomic spread of Cajas's paramo rodents (1 oryzomyine, 3 thomasomyine,
1 ichthyomyine, 2 akodontine, '. phyllotine, and 1 sigmnodontine), when compared
to rodent communities in Peru (e.g., Pearson and Ralph, 1978; Patton, 1986), does
not support Reig's (1986) contention that, while akodontines and phyllotines are
the most speciose myomorph rodents in puna habitats, the Oryzomyini are the most
speciose in paramo.



Caenolestes caniventer Anthony, 1921 and C. tatei Anthony, 1923.-On 9
September 1983, a pregnant female (1+1)"4 of Caenolestes caniventer (see Fig. 5)
was caught in one of 20 snap traps placed in shrubby secondary forest in the Rio
Mazan valley. Three days later, a specimen of C. tatei was taken in the same

14 i.e., one embryo in each uterine horn.

Figure 5. Caenolestes caniventer. Photograph, Adrian Barnett.


habitat type on a moss-covered branch some 2 m from the ground. Both species are
comparatively little-known. Such habitats are known to be favored by caenolestids
(see references in Barnett, 1991). Caenolestids can be locally common (Kirsch and
Waller, 1979). However, trapping in subsequent years on the Cajas Plateau found
no more specimens, in neither the primary nor secondary forest of Mazin, nor the
other montane forest site at in the Zorracucho Valley. Nor were they caught on the
pAramo, supporting the contention of Hunsaker (1972) that both these marsupial
species are exclusively montane forest species.
C. caniventer is widely distributed, extending from central Andean Ecuador
(Eisenberg and Redford, 1999) to northern Peru (Pacheco et al., 1995). C. tatei has
a much more localized distribution. Barnett (1991) has pointed out that the Rio
MazAn valley is very close to G.H.H. Tate's original collection site for C. tatei,
Molleturo, Azuay Province (see Anthony, 1921), and suggested that C. tatei may
be endemic to this massif. The recently described Caenolestes condorensis (Albuja
and Patterson, 1996) also shows this pattern of restricted distribution on a single
massif, being confined to the Cordillera del C6ndor, southeastern Ecuador.


Cryptotis montivaga Anthony, 1921.-Twelve specimens of C. montivaga
were trapped. One was taken in montane forest, five in quenoa forest, and six in
streamside scrub (including one in the pAramo/forest ecotone at Chirimanchi).
None was taken in open grassland habitats (see Barnett, 1992). These preferences
are similar to those recorded for C. thomasi (Durant and P6faur, 1984; Lopez-
Arevalo et al., 1993) and C. meridensis (Durant and Diaz, 1995). Head and body
length of trapped specimens of C. montivaga varied between 65 mm and 86 mm.
Weights varied between 9 g and 16 g, making C. montivaga one of the largest
members of the genus. This may be associated with the altitude at which it lives,
for it is the highest-living member of the genus, and metabolic constraints on
small-bodied insectivores are severe under the climatic conditions prevailing at
high altitudes (see Choate, 1970; Doucet and Bider, 1974; Pankakowski, 1979;
Churchfield, 1990; Barnett, 1992).
It is common for shrews to die of fright in live traps (Orr, 1949; Churchfield,
1990; Lopez-Arevalo et al., 1993). However, the montane forest specimen
(weighing 10 g when captured) was captured alive in a streamside live-trap set for
Chibchanomys. It was maintained in captivity for several days, allowing a unique
opportunity for observation (there being no other data on the behavior of this
species). It ate grasshoppers, crickets, staphylinind, chrysomelid and ruteline
beetles, muscid dipterans, hemipterans, hairless caterpillars, small moths
(including Pterophoridae, Pyralidae and Tortricidae), phasmids, and earthworms.
All were killed by a bite to the 'neck.' After initial investigations, it avoided
harvestmen (Phalangida), ants, and hairy caterpillars. It was unable to deal with
the heavy annour of scarabaenid beetles or larger members of the Rutelinae, and


became mired in the wing-scales of large moths. It ate up to 2 g of food in a two
hour period. As noted by Churchfield (1990), food-finding in many Soricidae
appears to be largely a matter of chance. These observations compliment the
preliminary data on the diet of C. montivaga presented by Barnett (1992), who
noted that the contents of five stomachs included beetle elytra, spider legs, and
Of the five females caught, two were lactating (NHM 84.338, 87.919: caught
on 29 August 1983 and 28 July 1984, respectively) and one (now in the collection
of the Esquela Polytechnica Nacional, Quito, trapped 20 August 1981; head and
body 82 mm, weight 16 g) was pregnant with 2 embryos (1+1). Lopez-Arevalo et
al. (1993) reported C. thomasi in Colombia breeds in these months. However, this
period represented the rainiest time at their eastern Cordillera study site, unlike
Cajas, where these are the driest months.
A preliminary report in Barnett (1992) on the litter size of C. montivaga has
now been supported by three additional observations. In each case the females had
two embryos. In Barnett (1992) I pointed out that C. montivaga's observed litter
size was low compared to the rest of the genus Cryptotis (see Choate, 1970; Mock
and Conway, 1975). Relating it to the rigours of insectivorous life at high altitude,
I ascribed the small litter-size to K-selection. I now believe this is more likely to be
due to adversity-selection, the form of investment-mediation in offspring that
operates in predictably severe environments (see Southwood, 1977), being (in the
case of C. montivaga) a consequence of lactational stress (see Sikes, 1995;
Rogowitz, 1996)1. This result and rationale runs counter to early studies (e.g.,
Dunmire, 1960) that suggested that litter size might increase with increasing
altitude as a compensation for a shorter period of reproductive activity.
Marshall (1980), pointing out that the extinction of the most specialized
caenolestids (of the sub-family Adberitinae) was contemporaneous with the
invasion of the Neotropics by placental insectivores, believes that ecological
similarity and competition was the cause of their demise. Extending this notion,
Barnett (1992) posited that the low numbers of C montivaga trapped in montane
forests at the Rio MazAn were due to scramble competition for insects with the
area's caenolestid marsupials (sub-family Caenolestinae). This is supported by
reports from wildlife filmmakers Jim and Teresa Clare (pers. comm.) that
Cryptotis is common in dense secondary forest around Hacienda Majan, near
Molleturo (2318 m). They reported that caenolestids are rare or absent from this
property. A similar habitat separation was recorded by Lopez-Arevalo et al. (1993)
for Cryptotis thomasi and Caenolestes obscurus.
C. montivaga was originally described by Anthony (1921) from specimens
collected at 3050 m near Besti6n, Azuay Province, Ecuador. When collected the
specimens from Cajas were the second known series. Skulls in owl pellets from
Podocarpus National Park, Loja (A. Barnett pers. obs.), indicate that the species
ranges farther south than both these sites. C. montivaga was formerly considered

15 this is consistent with the recent report by Woodman and Timm (1999) of 4 embryos in Cryptotis
goodwini magnimana, an animal known from forests at 1730 m.


the most southerly known Neotropical insectivore (see Barnett, 1992). However,
this mantle now rests with Cryptotis peruviensis, recently described by Vivar et al.
(1997) from northern Andean Peru.


Akodon mollis Thomas, 1894 and Akodon orophilus Osgood, 1913 -
Togeather these two species comprised 70.8% of all identified animals (342
specimens; 117 in montane forest, 225 in pAramo) and were numerically dominant
in all habitats types, except streams and quaking bogs. Robert Voss (1983, 1988,
pers. comm.) obtained similar results in northern Ecuador. Luis Albuja found
Akodon aerosus to be the most abundant small mammal of the Cordillera del
C6ndor (see Schulenberg and Awbrey, 1997: 73) and James Patton (pers. comm.,
1998) reports trapping 150 individuals with 35 museum special snap traps in one
36 hour period in less than one hectare of high-altitude bunch grass. In Cajas, most
piramo specimens (208, 92.4%) were taken in cover (quebradas, streamsides, and
quenoa), and only 17 (7.6%) were taken out in the grasslands of the pAramo. Dorst
(1958) found that both Akodon (now Chroeomys) jelskii and A. boliviensis also
preferred sheltered habitats.
All specimens from Cajas were originally assigned to a single species, A.
mollis. However, following comments from James Patton (pers. comm.), specimens
were re-examined, revealing the presence of several specimens of A. orophilus in
the collection. Formerly considered a subspecies of A. mollis, this taxon is now
considered a full species (Myers et al., 1990; Patton and Smith, 1992). The
presence of A. orophilus in Cajas is a new species record for Ecuador and
constitutes a range extention of some 250 km from the species' previously known
northerly limit in northern Peru (see Cabrera, 1961; Myers et al., 1990; Patton and
Smith, 1992; Wilson and Reeder, 1993).
The ecology of the genus Akodon appears to closely resemble that of the
temperate genus Microtus, to which, superficially, Akodon bears a strong external
resemblance (see Fig. 6). Several members of the genus have been well studied (see
references below), but both A. mollis and A. orophilus appear to be little known.
From data in Cabrera (1961), Nowak (1999), Eisenberg and Redford (1999), and
examination of the NHM collection, A. mollis would appear to be one of the
highest-living members of the genus. Specimens varied from 75 mm-118 mm in
head and body length. The tail is always shorter than the head and body (63 mm-
88 mm). Weight varied from 15.5 g for sub-adult males to 43 g for a pregnant
An analysis of fecal pellets from 17 A. mollis and 6 A. orophilus from Cajas
across a range of head and body lengths found both plant and insect remains in the
diet. Insect remains varied from 10% to 70% by volume per individual. These
figures closely resemble those obtained by Barlow (1969) for A. azerae, where an


Figure 6. Akodon mollis. Photograph, Adrian Barnett.


analysis of 11 stomachs found 70% invertebrates and 20% plant material (see also
Eisenberg and Redford, 1999).
This dietary data, combined with ubiquity and high numbers in all habitats
(see Table 3) indicate that, in Cajas, A. mollis is a widespread and abundant
generalist. Such an ecology resembles that of other akodontines (Chroeomys
andinus in Peru, Pearson and Ralph, 1978; A. azarae in Argentina, Dalby, 1975;
Apfelbaum and Blanco, 1985a; Ellis et al., 1997; A. boliviensis in Peru, Dorst,
1958; A. delores in Argentina, Apfelbaum and Blanco, 1985b; Piantanida, 1987;
Abrothrix olivaceous in Chile, Fulk, 1975; Mur(ta and GonzAlez, 1985; Simonetti
et al., 1985; Meserve and Le Boulenge, 1987; Muriua et al., 1987; Abrothrix
longpilis in northern Chile; Glanz, 1984).'6 A. orophilus, by contrast, appears to
occur at comparatively low densities in closed cloud forest habitats.
The two species appear to show ecological separation. All specimens taken in
in the Podocarpus forests at MazAn were A. orophilus, while those in the adjacent
grazed grassland were A. mollis. In the Zorracucho Valley, A. orophilus was taken
in the secondary forest on the sides of the valley while A. mollis occupied the
grazed grasslands of the valley floor. However, A. orophilus was not caught in
pAramo, and there all the akodons caught in quenoa forest were A. mollis,
indicating it is another factor (possibly temperature) rather than cover that limits
the upper altitudinal range of A. orophilus."
Species of Akodon have been variously reported as diurnal, nocturnal,
crepuscular or active at any time of day (Eisenberg, 1989; Emmons, 1997; Nowak,
1999; Eisenberg and Redford, 1999). A. mollis appears to be predominantly
nocturnal. Of 225 individual A. mollis trapped on the pAramo, only nine (4.0%)
were found during mid-day trap checks.
Reproductive data does not appear to have been recorded before for A. mollis
nor for A. orophilus. No pregnant A. orophilus were captured, but three A. mollis
females were pregnant when trapped. Each carried two embryos (1+1). Other
members of the genus usually have three to four young (Nowak, 1999; Eisenberg
and Redford, 1999). As with Cryptotis, this difference may be attributable to
adversity selection (Southwood, 1977), a consequence of life at high altitudes (see
also Badyaev, 1997). A specimen of A. mollis collected in Loja (3 June 1899,
unaccessioned NMH specimen), was recorded by its collector, P. 0. Simons, as
having four embryos. Significantly, the collection altitude (2100 m) is much lower
than for any of the pregnant Cajas specimens, supporting the adversity selection
position. A study by Krohne (1980) on Microtus californicus populations under
conditions of distinctly different adversity (perennial native grasslands and
introduced annual grasslands), provides a supportive parallel to the Akodon results.
From Southwood's (1977) model one would predict that A. orophilus would have

16several of these were considered to belong to the genus Akodon when the papers in question were written
(e.g., Ab. olivaceus and Ab. longipilis, see Spotorno et al., 1990; Chroeomys andinus, see Patton and
Smith, 1992).
17a species whose fur is both shorter and less dense than that of A. mollis.


larger litters than A. mollis. It is to be lamented, therefore, that no reproductive
data is available from A. orophilus in Cajas to test this assertion at a local level.
Overall, 80% of Akodon captured in Cajas were juveniles (76.7% for A.
mollis and 83.3% for A.orophilus). Most (70%) of the adults collected were caught
in the last third of the fieldwork season. This suggests that A. mollis may have two
litters a year and a bimodal breeding season. This reproductive pattern has also
been reported for a number of other members of the genus (Nowak, 1999). It is
possible there is little temporal overlap between reproductive generations of A.
mollis (as also reported for Akodon spp. by Meserve et al., 1990) so that, during
August-September, the Cajas population is entering a second reproductive period,
with juveniles being a result of a preceding reproductive peak (see also Zuleta and
Bilenca, 1992). Dalby (1975) reported the average gestation time of A. azarae to be
22.7 days and the time to first breeding to be 2 months. If the time-scales of A.
mollis are similar, this would place the other reproductive peak around May-June,
the time of highest rainfall on the Plateau (see Meteorological Office, 1977;
Ramsay, 1992). On a number of other Andean pAramos this has been shown to be
the period of greatest bio-productivity (Ramsay, in press, Ramsay and Oxley, in
press), reflecting peaks in both fresh plant growth (Sturm and Rangel, 1985;
Ramsay and Oxley, 1996), and insect abundance (Sturm, 1979; P. Ramsay pers.
comm., 1998).
Osgood (1914: 164) noted that grassland populations of A. m. orophilus 18
resembled Microtus with "labyrinthine runways, open burrows, and fresh grass
cuttings," while "in heavy woods or rocky stream beds they] lead wandering lives,
and have as retreats only natural openings." Data from the current study supports
this observation. No akodonts were trapped from burrows in montane forest or in
quenoa forest. However, on the paramo, they were trapped on several occasions
emerging from holes. These frequently had small mounds of fresh vegetation and
grass seed heads piled next to them. The burrow entrances often occurred in
aggregated groups of a dozen or so. During the day most would be sealed with a
cap of earth. Dorst (1958) observed cut vegetation by and within the burrows of A.
The importance of melanic variation in pelage coloration in A. mollis has
been difficult to determine. Writing of color variation in A. mollis, Hershkovitz
(1940a: 1) noted that "despite the diversity of their habitats, populations of this
mouse exist practically undifferentiated [from] the inter-Andean Plateau ... nearly
[to] the snow line." There was concordingly little variation in specimens from
Cajas, variation being limited to some animals with a wide, ill-defined dorsal
stripe. However, four very dark specimens were taken at Yakatuviana (NHM
82.808, 82.809, 84.292, 84.293). Partial melanism is often associated with
mammal populations living in moister warmer environments such as occur at
Yakatuviana. Osgood (1914), for example, named A. mollis orientalis 1" from dark-

1now considered a sub-species ofAkodon orophilus (see Patton and Smith, 1992 page 91).
19now considered a sub-species ofAkodon orophilus (see Patton and Smith, 1992 page 91).


furred specimens from the humid montane forests of north-eastern Peru, and
Sanborn (1947) noted that inambarii, a dark subspecies of Peruvian and Bolivian
Chroeomysjelskii (formerly Akodon jelskii), came from an area that is notably wet
and cloudy.20 However, examination of 244 Ecuadorian A. mollis skins in the
NHM collection revealed very few other melanics or partial melanics. Examination
of another 261 specimens from 14 other Akodon species revealed no within-species
pattern of pelage color variation that could be associated with habitat.21 Like the
NHM's leucystic specimen of A. olivaceus in the same collection (NHM,
the specimens from Yakatuviana should perhaps be best considered sports and un-
deserving of subspecific status (see Owen and Shackelford, 1942; Mustrangi, 1994;
Literik and Zejda, 1995 for similar examples with other species). The trapping of
several similarly colored animals in the same locality may simply indicate genetic
relatedness and a sharing of the same colour-coding allele (see Benton, 1953;
Fedyk and Borowski, 1980 for parallel mammalian examples).
Of the 168 A. mollis skins presented to the NHM, 19 had white patches of fur
about the head and neck (1.2%). The causes) of the white fur patches are
unknown. They were constant in neither size, shape, nor position. Field dissections
revealed neither subcutaneous parasites at these locations, nor any scars or lesions.
However, white spotting is frequent in European shrews (Sorex) (see Pugek, 1964),
where it is frequently a result of intra-specific aggression, usually between males
(Churchfield, 1990). Fifteen of the A. mollis with white spots were adult (78.9%, 6
males, 9 females), indicating that a similar explanation, combined with aggression
during mating, may apply here.
Chibchanomys orcesi Jenkins and Barnett, 1997.-Five specimens of this
new species were captured (see Fig. 7). Two were taken on a small bush-covered
island in the middle of a small, cold, clear, fast-flowing stream at 3700 m. Flowing
between large boulders, the stream had a bottom of course of gravel and stones. It
was no more than 40 cm deep and lacked aquatic macrophytes (see Fig. 8). Two
more were caught in similar habitat at Lake Torreadora, 4000 in. A fifth specimen
was caught in the Zorracucho Valley, in a broad, deep muddy stream feeding in to
Lake Llaviuco. Heavy rain may have washed the animal in from higher pairamo
streams. Further details on habitat are given by Barnett (1997a).
No ichthyomyines are specialized for lentic waters, though Ludovic
Soderstrom caught Ichthyomys in marshy fields (Tate, 1931), and local informants
told Andrea Pogson (pers. comm.) that they had occasionally seen fishing mice
(presumably C. orcesi) in lakes on the Cajas Plateau. The use of mid-stream
islands seems a common feature of aquatic mice. Described as successful trapping
sites for many ichthyomyines (see Voss, 1988; Barnett, 1997a), they have also been

20 for other examples see Nowak, 1991: 658-659 on the Himalayan vole Hyperacrius; Rosevear, 1969,
1974; Barnett and Prangley, 1997 for examples with African mammals.
21 though, as J. Patton (pers. comm.) has pointed out, all of the Andean wet forest Akodon (aerosus, fumeus,
kofordi, mimus, orophilus, siberiae, surdus, torques) are dark in comparison to open grassland species
boliviensiss, Juainensts, lutescieus, mollis, subfuscus).

Figure 7. Chibchanomys orcesi. Photograph, Jim Clare.


Figure 8. Habitat shot for C. orcesi. Near Lake Torreadora (4000 m).Photograph, Adrian Barnett.


found to be so for their ecological equivalents in the Paleotropics (e.g., see Osgood,
This new species (Jenkins and Barnett, 1997) was named in honor of Gustavo
Orc6s, a pioneer of Ecuadorian vertebrate biology.22 Its ecology, natural history,
and biogeography were discussed by Barnett (1997a). Like Caenolestes tatei, it is
believed to be endemic to the Cajas Plateau (Barnett, 1997a). It is the sixteenth
known ichthyomyine (see Voss, 1988; plus Ochoa and Soriano, 1991) and the sixth
member of its tribe known from Ecuador, making that country the most speciose
for this tribe.
In his review of the Ichthyomyini, Voss (1988) gives the distribution of the
other member of the genus, C. trichotis, as the highlands of the Tachira Andes,
Venezuela, the Colombian Cordillera Oriental, and the Cordillera Carpish in Peru.
C. orcesi appears to occur at just one locality within the range of these widely
separated populations. Barnett (1997a) provided an analysis with examples of
parallel biogeographical patterns in other taxa, including birds, small mammals,
and bromeliads. However, Paulina Jenkins used the PAUP program (Swofford,
1990) to compare the features shared by C. orcesi, the Peruvian "C. trichotis," and
Colombian-Venezuelan populations of C. trichotis. Her data reveal that the
Peruvian population may, in fact, be more closely related to C. orcesi (see Jenkins
and Barnett, 1997). Though this pattern does not occur in all taxa (see Pearson,
1982; Reig, 1986; Albuja and Patterson, 1996; Vivar et al., 1997), this Peruvian-
Ecuadorian/Colombian-Venezuelan distribution pattern is also far from unusual in
high altitude vertebrates, with the Huancabamba Depression acting as a
distributional barrier (see Duellman, 1979; FjeldsA and Krabbe, 1990 for parallel
herpetological and avian examples, respectively)23.
Stomach contents of three specimens showed fish and larvae of aquatic
insects. Observations of a captive specimen by J. and T. Clare revealed the animal
could actively hunt for fish underwater. The small eyes and enlarged vibrissae
make tactile location the most feasible form of hunting. The ecology of this species,
like most of the tribe, closely resembles that of the Old World otter shrews
(Micropotomogale lamottei, M. ruwenzorii, and Potomogale velox; see Kingdon,
1997, for an account). The African lowland marine Colomys goslingi also shows
similarities in feeding ecology (Dieterlen and Statzner, 1981), though it is less
morphologically specialized (Kingdon, 1997).
C. orcesi appears to be quite rare. Persistent trapping in appropriate localities
over several years by two separate study groups yielded very few specimens (see
Barnett, 1997a). In each case animals were trapped from the same localities, and
not from other, apparently suitable ones. The species therefore appears to have both
low population density and restricted habitat preferences. Also, it is only known
from the Cajas Plateau (Barnett, 1997a; Jenkins and Barnett, 1997). Together these
factors provide cause for conservation concern. Small riparian mammals are

22 it is the second higher vertebrate named in his honor, see Ridgely and Robbins (1988).
23 Vivar et al. (1997) believe that this 'pattern' may be (partly) explained by sampling bias, reflecting the
paucity of work in this border region.


frequently at risk from anthropogenic habitat changes (R.G. Anthony et al., 1987;
Galindo-Leal, 1997). Barnett (1997a) has cited reasons why C. orcesi populations
in Cajas may be threatened. These include burning, grazing and increased tourist
pressure. However, such concerns should not be given uncritically. Louise Emmons
(pers. comm., 1997) has received reports of the Central American ichthyomyine
Rheomys being observed in quite polluted water, and Tate (1931) reported
Neusticomys from irrigation ditches.
Microryzomys altissimus (Osgood, 1933) and M. minutus Tomes, 1860.-
Thirty M. altissimus and two M. minutus were trapped. The relative numbers may
have little overall significance, for Durant and Diaz (1995) have reported that M
minutus numbers show great seasonal variation. Along with Thomasomys gracilis
(see below), M. minutus is the smallest rodent on the Plateau, with individuals
being recorded from 9.5 g. M. minutus was taken in secondary forests around Lake
Llaviuco in the Zorracucho valley (3100 m). M. altissimus was also found below
the tree-line (two records from Zorracucho) and in the shrubby forest/pAramo
ecotone trapped at Chirimanchi, but was most common in the pAramo. On the
paramo most records come from sheltered habitats (rocky outcrops, 9; Polylepis
forests, 7; streamside scrub, 6; quebradas, 3). The animal was infrequent on the
exposed pAramo grasslands (3 of 28 records). This preference for sheltered habitats
is shared by the shrew Cryptotis montivaga (see above) and pAramo-living species
of Caenolestes (see Gregory, 1922; Tate, 1932a).
Based on museum specimens, Carleton and Musser (1989) have noted that
the range of M. altissimus extends to higher altitudes (to 4500 m), and M. minutus
favors altitudes between 2000 and 3500 m. Handley (1976) and Osgood (1933)
both record M. minutus almost exclusively from montane and cloud forests (the
latter as aurillus). Where M. minutus does enter the paramo zone, it is restricted to
heavy cover such as is offered by Polylepis and Espeletia thickets (the field notes
of R. Voss, quoted by Carleton and Musser, 1989: 56-57). Based on analysis of
over 900 museum specimens from 200 collecting localities, they report that 75% of
the records for minutus are concentrated between 2000 and 3500 m, while 80% of
the records of altissimus occur between 2500 and 4000 m. They record that a zone
of sympathy exists between the two species near the tree-line. At 3100 m, the
records from Zorracucho are in agreement with this synopsis.
The two species are difficult to separate, both in the field and in the museum.
However, field experience has shown that, while minutus generally has a
monocoloured tail, altissimus has one which is bicoloured, dark above (see Barnett
and Muleon, 1999). This has also been observed in museum material by Carleton
and Musser (1989), who have also made helpful observations on the differences in
shape and arrangement of the planar pads and in the shape of the hind foot,
characters that may also be used to distinguish these two species in the hand.
Carleton and Musser (1989) also report the tail to be proportionatly longer (145%
vs. 137% of lib) in M. minutus than in M altissimus. Eisenberg and Redford
(1999) provide characters of the palate for distinguishing skulls of the two species.
Fifteen of the M. altissimus specimens were immature. One male (NHM
82.732, caught 7 August 1983) had scrotal testes, one female had a perforate


hymen (caught 16 July 1983). Neither of the M. minutus were in breeding
Oligoryzomys destructor (Tschudi, 1844).-Five specimens (3 males, 2
females) were trapped. All specimens came from the Zorracucho Valley (3100 m).
Other than an avoidance of grasslands trapping data gives little indication of
habitat preference by this species. One individual (NHM 84.300) was caught on the
mixed mud and gravel banks of the same unnamed stream that yielded
Chibchanomys orcesi (see Barnett, 1997a). An adult female (NHM 84.303) was
living in human habitation. One juvenile male (NHM 84.299) was taken on an
open disturbed wooded slope with sparse ground cover, no streams and dry exposed
soil, while another (NHM 84.302) was taken on a stream-rich slope with thick
vegetation and mossy groundcover beneath scattered Eucalyptus globulus.
Examination of labels of specimens in the NHM collection shows a similar
variability in recorded habitat preferences throughout the collection, with
references to forest (NHM, a hedge (NHM, a house (NHM, streamside (NHM, and swampy grassland (NHM Most Ecuadorian 0. destructor specimens in the museum collection
do not come from altitudes as high as those at Zorracucho, being more commonly
taken around 2900 ft (= 1000 m). Hershkovitz (1940b) recorded this species (as 0.
spodiurus) from 1500 m in the sub-tropical forests of Ecuador's western Cordillera.
Osgood (1914) reported it (as 0. stolzmanni maranonicus) from dense humid
forests and town gardens in north-eastern Peru.
Oryzomys albigularis Tomes, 1860.-Sixteen 0. albigularis were obtained.
This beautiful rodent is one of the largest on the plateau, weighing up to 79 g with
a head and body length of up to 162 mm and a tail length of 170 mm. All belong to
the subspecies albigularis. Members of this subspecies are distinguished from 0. a.
moerax by their belly colour (paler in albigularis) and the presence of a throat
patch where the hairs are pure white to the base (a feature absent in 0. a. moerax)24
(see Gyldenstolpe, 1932: 14). On Cajas specimens of 0. albigularis were trapped
only in cloud forest, a habitat preference also reported by Osgood (1914), Handley
(1976), Ashe and Timm (1987), and Eisenberg and Redford (1999). It appears to
prefer dense cover (Osgood, 1914). Durant and Diaz (1995) reported that this
species was at its lowest density in the wet season. Such seasonal fluctuations may
explain why, while P6faur and Diaz-de-PAscual (1985) reported 0. albigularis as
the numerically dominant species at northwest facing montane forest sites between
1000 and 2500 m, it was the rarest species in Zufiinga et al. 's (1983) study.
Six of ten males were of adult size, four were in breeding condition (NHM
82.725, 82.276, 84.306, 84.308). A female (NHM 84.305), taken on 3 September
1983 was lactating. Two of the specimens were retrieved from a number of 0.
albigularis found dead on a path at Yakatuviana. Though suffering some insect
damage, all were otherwise unmarked. No reason could be found for this
phenomenon. Interestingly enough, in the original description of the species Tomes

24 NHM specimens examined: 54. 413 to 417, 54.419,


(1860: 265) noted that the holotype and topotype were both found under similar
circumstances, lying dead on a [forest] path among a number of other of dead
conspecifics. Examination of these specimens (inc. NHM shows they
also lack obvious external damage. None of the females trapped in Cajas were
pregnant. Timm et al. (1992) have recorded litter size in this species as three.
It has long been known that this species had a species of staphylinind beetle
Amblyopinus in its fur (e.g., Seevers, 1950). This association has been shown
(Ashe and Timm, 1987; Timm and Ashe, 1987) to be commensal rather than
parasitic, with the beetle feeding on such ectoparasites as mites and fleas in the
rodents fur. The number of beetles has recently been shown to vary between sexes
and ages of these rodents (Barnett, 1997b; Barnett and Shapley, in prep. b)25 in
accordance with predictions of theories of androgen-mediated immuno-suppression
and sex-bias in mammalian ectoparasite loads (Schalk and Forbes, 1997). As far as
we know, this is the first time such an indirect effect has been shown.
Phyllotis andium Thomas, 1912 and Phyllotis haggardi Thomas, 1908.-
These are the most northerly of the 11 species in the genus Phyllotis (Pearson,
1958, 1982; Braun, 1993; Nowak, 1999). One P. andium and six specimens of P.
haggardi were taken. The two species are easily separated in the hand (Barnett and
Muleon, 1999). P. haggardi has a bicoloured tail that is usually greater than 95
mm in adults, whereas P. andium has a unicoloured tail that is usually less than 85
mm. Only one female P. haggardi, an immature, was trapped (NHM 82.811).
Three of the five males captured were mature and in breeding condition (NHM
82.812, 84.344, 84.345).
One P. haggardi, a young male, was found living commensally in the
warden's hut at Lake Torreadora. All other specimens were taken in or around
rocky outcrops or stony ridges in otherwise open pAramo where the shrub cover
was limited. All captures were made at night. Though taken at the same altitude,
the two species were not sympatric. There were substantial habitat differences, with
the single P. andium taken from amongst Hieracium and Baccharis bushes in a
shallow shrub-rich valley, while P. haggardi occurred in rockier more exposed
habitats. In Peru, Dorst (1958) found similar habitat separation occurring between
(respectively) Phyllotis pictus and P. darwini. According to Pearson (1958, 1972),
P. andium inhabits brushy habitats, but neither he, Anthony (1924a), Braun
(1993), nor Eisenberg and Redford (1999) give any indication of habitat
preferences for P. haggardi.
P. haggardi was previously known only from north-central Ecuador (Wilson
and Reeder, 1993; Eisenberg and Redford, 1999),26 making the records from Cajas
range extention for the species of several hundred kilometers.
Cajas is unusual in the phyllotine component of its small mammal
community. P. haggardi is endemic to Ecuador and is at its southern limit in
Cajas, while P. andium is close to its northern limit there (see Pearson, 1972;

25 manuscipt available from author on request.
26 contra Pearson (1972) and Braun (1993) who believed it to occur in southern Ecuador also.


Braun, 1993). The zone of overlap between the two species is only some 110-160
km long and is further restricted by altitude; while P. andium ranges widely from
200 to 4800 m, P. haggardi occurs only between 2000 to 4500 m (see Pearson,
1958, 1972; Braun, 1993: table 1). This may account for Pearson's (1982: 273)
opinion, now obviously erroneous, that the two species do not occur in the same
geographical location. According to Pearson (1958: 441), in P. haggardi there is a
trend of "increasing duskiness to the south, together with an increased length of the
skull and lengthening of the nasals" (see also Anthony, 1924a). He uses the
appellation P. h. fuscus for southern populations, including those in the Cajas
Sigmodon inopinatus Anthony, 1924.-Five specimens of S. inopinatus were
trapped. All were taken near water; one in a hole in a riverbank quebrada at
Chaupiurcu (3750 m), another in a marshy area at Lake Torreadora (4000 m), and
three others on a dense reed mat forming a quaking bog in the middle of a shallow
unnamed lake near Lake Torreadora. Surface runs were wide and obvious at the
reed mat trap site. Similar habitat preferences and run-making behaviour have
been reported for S. hispidus (e.g., Mazzotti et al., 1981; Fitzgerald, 1983) and S.
alstoni (Voss, 1992). No nests were seen.
One female specimen (now in the collection of the Escuela Polytechnica
Nacional, Quito) taken on 19 July 1984 had four embryos (2+2). This is at the
lowest end of litter size variation in Sigmodon (see Randolph et al., 1977;
Mattingly and McClure, 1982) and, since S. inopinatus is the highest-living
member of the genus (see Voss, 1992), may be attributable to the same adversity-
selected suite of traits also believed to be governing small litter size in Cryptotis
and Akodon in Cajas (see Southwood, 1977). In the only other datum point for this
species, a litter of four is also reported by Anthony (1924b) from the type specimen
of the species, the largest individual in a series of 11 specimens. Nipple number is
lower in S. inopinatus than in other members of the genus (Voss, 1992).
S. inopinatus shares many aspects of the ecology of its close relative S.
hispidus (see Voss, 1992). However, while S. hispidus is the most abundant
member of the small mammal community in many parts of its wide range (Sietman
et al., 1994; Doonan and Slade, 1995; Stokes, 1995), relatively low numbers of S.
inopinatus were found at Cajas. This may be due to competition with the
numerically dominant Akodon mollis. Though smaller, A. mollis appear to share a
similar ecology to S. inopinatus and may have excluded it from some habitats.
Such interactions have been recorded at various locations in the extensive range of
the genus Sigmodon (Glass and Slade, 1980; Putera and Grant, 1985; Davis and
Ward, 1988; Voss, 1992). Such interactions may be especially important in Cajas
where populations of S. inopinatus would appear to be at the southern limits of
both the species' and genus' range (see Voss, 1992)27 and hence possibly a less
effective competitor (see Grant, 1972, who includes a discussion of competitive
exclusion effects at range limits in rodents).

27 Only S. peruanus, which extends into northern Peru, occurs further south (Pacheco et al., 1995; V.
Pacheco pers. comm., 1999).



The records of S. inopinatus at Cajas challenge an evolutionary scenario for
the species proposed by Reig (1986). The type series of S. inopinatus was taken by
G.H.H. Tate in October 1923 at Urbina at 11,400 ft (3508 m) on the slopes of Mt.
Chimborazo (Anthony, 1924b). The Cajas specimen series is the second ever for
this species (Voss, 1992) and represent a southerly range extension of around 200
km. According to Reig (1986), the genus arose in north-central Peru from a
Neotomys-like ancestor and moved northwards to its current distribution, via
Andean valleys. Reig (1986: 430) proposes that S. inopinatus is a relictual and
historically isolated species that "is likely to have arrived [on the pAramos of
Chimborazo] by the uplifting of the pAramos during Plio-Pleistocene times,"
presumably while this northward migration of the genus was in progress. With the
discovery of the species at Cajas, such a site-specific explanation is clearly no
longer required, especially since it is likely that further trapping will find the
species at other suitable habitats between the two currently known localities.
Though named as a full species by Anthony (1924b), this taxon was (with the
exception of Reig [1986] who accepted its original status), generally considered a
subspecies of Sigmodon hispidus until Voss' revision of the genus (Voss, 1992).
Noting that the long silvery rump hairs were not diagnostic as Anthony (1924b)
proposed, Voss (1992) provides a series of cranial, dental, and chromosomal
characteristics to support his re-elevation of this taxon to specific status (contra
Hershkovitz, 1955; Cabrera, 1961). The parapatric S. peruanus, also recorded from
Azuay Province, is found only in lower altitudes and more xeric habitats (see Voss,
Thomasomys spp.-Six species of Thomasomys were taken: T. aureus
(Tomes, 1860), T. baeops (Thomas, 1899), T. cinnameus Anthony, 1924, T.
gracilis (Thomas, 1917), T. paramorum Thomas, 1898, and T. pyrrhonotus
Thomas, 1886. Only T. cinnameus occurred in pAramo alone. T. pyrrhonotus and
T. gracilis were taken in pAramo and montane forest. The other three species were
taken only in montane forest.
The paramo-living species of Thomasomys appear to favor sheltered areas.
Based on trappability, Anthony (1924b) believed T. cinnameus to be rather rare.
Instead, results from Cajas indicate it is a quenoa forest habitat specialist. In the
current study, four of the six T. cinnameus were caught in minimally disturbed
quenoa forests, all on the ground in areas with dense canopy and little groundcover
but deep moss. The two other specimens were taken in a dense isolated clump of
composite bushes on an exposed slope covered with tussock grass. It was the only
member of the genus to be found solely in pAramo.
Three specimens of T. pyrrhonotus were taken in degraded quenoa forests
with open canopies and heavy shrub layers. Two T. pyrrhonotus specimens from
montane forests were also taken in degraded vegetation (one on an open wooded
slope, the other among rocks in a grass-covered area logged some five years
previously). Osgood (1914) also caught this species both above the timberline and
in cloud forest.


Of 25 T. gracilis trapped, 21 (74.4%) were from habitats in the piramo.
Though several T. gracilis were trapped in sheltered areas such as dense scrub,
streamsides, quebradas, and rock jumbles in piramo grassland, as well as in
quenoa forests, none was taken in open, exposed piramo grassland. Some were
taken in degraded, open-canopied quenoa forest at Chaupiurcu, but only in the
most sheltered parts. Both T. pyrrhonotus and T. gracilis were trapped in the lower
limbs of Polylepis trees. Nowak (1999) indicates arboreality is common in this
genus. However, otherwise terrestrial species can be attracted to baited traps set in
trees (Manville et al., 1992). Such confounding effects, combined with the jumbled
and topographically complex nature of this forest type (see Fig. 3), means that this
may not be a true indication of regular arboreal activity for these species.
Specimens of T. gracilis were also captured in secondary cloud forest around Lake
Llaviuco in the Zorracucho Valley, where it preferred mossy-floored habitats with
well-developed canopy and thick epiphyte cover.
Four T. aureus (see Fig. 9) were captured beneath dense shrub cover in
secondary forest between 2700 m and 3100 m, another on a moss-covered log deep
in primary montane forest at 2800 m, and two in very dense grass and reeds by
Lake Llaviuco (3100 m). Though records from Cajas, Colombia's Reserva
Biol6gica Carpanta (Lopez-Arevalo et al., 1993), and all specimens in the NHM
are from forested areas (see also Eisenberg and Redford, 1999), Reig (1986)
reported that this species may also occur in phramo. As in the current study, T.
aureus was infrequently trapped by Lopez-Arevalo et al. (1993), where it
represented less than 1% of the catch, indicating that it is either rare or difficult to
trap. So little is known of the ecology of this species that I consider it worth
recording that fieldnotes by Gilbert Hammond accompanying his specimens of T.
aureus (NHM,, report that the species nests in
trees and feeds on fruit of the grenadilla (any of several species of shrubby montane
forest Passiflora, Passifloraceae).
The 13 trapped T. paramorum came from a variety of habitats, including
secondary montane forest, bamboo thickets, and scrub on recently deforested land
but not (contra Reig, 1986) on the pAramo. The common denominator for all
chosen habitats seems to be moist mossy ground cover, fallen logs, and proximity
to stands of Chusquea bamboo.
All five specimens of T. baeops were trapped in disturbed secondary forest
sections of primary-secondary cloud forest mosaics. Preferred habitats had a dense
cover of shrubs, but little arborescent canopy development. Reig (1986) listed this
species solely as a pAramo-dweller.
A female T. gracilis (NHM 82.818, head and body 103 mm, 41 g) was
pregnant (1+1) when trapped on 29 July, 1981, another (NHM 84.339, head and
body 86 mm, wt. 16.25 g) was lactating when caught on 21 August, 1983. Males in
reproductive condition were captured between 15 and 28 August over three years.
Juveniles of T. gracilis numbered 15 (10 f, 5 m) (based on body size, pelage

Figure 9. Thomasomys aureus. Photograph, Vaughan Fleming.


grayness and pedal characteristics, see Barnett and da Cunha, 1998b). All juvenile
T. gracilis were between 64 and 100 mm (mean 83.06) in head and body length
(adult head and body mean = 98.0; range 86-116). Only one newly independent
individual was captured (head and body 64 mm, wt 10 g). No other trapped
Thomasomys females were pregnant. Two male T. aureus (NHM 84.324 and 25;
head and body 170 mm and 160 mm, weights 110 g and 102 g, respectively) had
scrotal testes (trapped 11 September 1983). One T. baeops (NHM 84.330; 104 mm,
43.5 g, trapped 4 September 1983) was lactating. One T. cinnameus (NHM 84.339;
head and body 86 mm, w 16.25 g) was lactating when trapped on 21 August 1983,
while a male (NHM 84.343; head and body 98 mm, weight 18.5 g) had scrotal
testes when trapped on 28 August 1983.
Habitat distribution data indicates a size/habitat pairing occurs amongst the
Thomasomys of Cajas. Each major habitat type (cloud forest and quenoa
forest/piramo grassland) having one large and two small species: T. aureus, T.
baeops, and T. pramorum for cloud forests, T. cinnameus, T. gracilis, and T.
pyrrhonotus for the pAramo. Head and body length has been proposed as a valid
measure of ecological separation between coexisting small mammals (see
Flowerdew, 1976). Size ratios for cohabiting pairs of Thomasomys is 1.43 for T.
baeops/T. pyrrhonotus and 1.57 for T. aureusIT. paramorum.8 This is close to the
ratios of 1.20-1.40 first noted by Hutchinson (1959) for morphological character
displacement among coexisting congeners. In each case the two smaller species are
separated by habitat preference, and one large and one small species inhabit a
closed-canopy arborescent habitat. Such mechanisms have been shown to structure
other Neotropical small mammal communities (Fulk, 1975; Meserve et al., 1982;
Vasquez, 1996) though such relationships are often complex (e.g., Meserve, 1981;
Murfa and Gonzalez, 1982; Ebensperger and Simonetti, 1996) and further work is
required in Cajas to analyze the fine details of the interactions of the area's small

Missing Species

Though clearly indicating a speciose locality, there are some puzzling gaps in
the current list of small mammals for Cajas. These may be due to a variety of
factors. As with bats (Koopman, 1978), birds (FjeldsA and Krabbe, 1990), and
amphibians and reptiles (Duellman, 1979; Lynch, 1979; Traub, 1979), the
dispersal barriers of the inter-Andean basins explain the absence in Cajas of some
species (e.g., Aepomys lugens is reported up to 3500 m, but only from the eastern
Andes). The absence of others known from the western Cordillera is most likely
attributable to altitude (e.g., Ichthyomys hydrobates to 2800 m and Oryzomys
auriventer to 1500 m).

28 metric data available from author on request.


There are a number of species whose absence cannot be explained
biogeographically, but for which possible ecological explanations exist. The semi-
aquatic ichthyomyine Anotomys leader has been recorded from Mt. Pichincha in
northern Ecuador to central Peru (Gardner, 1971). Its absence from Cajas may be
due to exclusion through competition with Chibchanomys orcesi, which may be
endemic to the Plateau (see Barnett, 1997a; Jenkins and Barnett, 1997). The
altitudinal and habitat preferences of these two species overlap (see Thomas, 1906,
Voss, 1988, for A. leader; Barnett, 1997a, Jenkins and Barnett, 1997, for C.
orcesi). At lower altitudes, members of the tribe Ichthyomyini have been recorded
in sympatry, with different species focusing on different food sources. Analysis of
data in Voss (1988) shows that, in such circumstances, one species is often
insectivorous, while another may have a high proportion of crab in the diet. A.
leader and C. orcesi both occur at the maximum recorded altitudinal elevation for
ichthyomyines (Barnett, 1997a), and at such heights freshwater crabs are lacking
(see data in Barnett, 1997a). Consequently the riparian habitats of Cajas's piramo
may be able to support no more than one species of this group.
On biogeographical grounds (see Gyldenstolpe, 1932; Cabrera, 1961;
Eisenberg and Redford, 1999), three other species of rodent, Rhipidomys
latimanus, Thomasomys silvestris and T. cinereiventer should occur in Cajas, but
have not been found there to date. Intense efforts were made to locate R. latimanus,
with targeted trapping in both cloud and quenoa forests for this arboreal species
(see Lopez-Arevalo et al.1993). Also, as noted previously by Barnett (1991), it is
unclear why the marsupial Caenolestesfuliginosus was not taken in paramo traps.
It is known to inhabit piramo and quenoa, and has been recorded from northern
Ecuador to northwestern Peru (Tate, 1931; Albuja and Patterson, 1996), where it
can be quite common.29
It should be noted that the Mountain Paca Agouti taczanowskii has not been
recorded in Cajas, though there are records at altitudes up to 3500 m for the
species to the south at Podocarpus National Park, Loja, where individuals visit
human habitations (A. Barnett, M. Prangley, and B. Larssen, pers. obs.; see also
Emmons, 1997) and in similar habitats in Colombia (Lopez-Arevalo et al., 1993).

Threats to the Ecology of the Cajas Plateau:

Alien species.-Mus musculus and Rattus rattus were also recorded. Both
were trapped in buildings, Mus in the warden's hut at Lake Torreadora and at an
unused brewery near Lake Llaviuco and Rattus at the trout farm at the Llaviuco
lakeside, where it was reported to be present in large numbers and feeding on
stored bags of fish feed. Conversations with the area's older residents revealed that
Mus and Rattus are probably recent arrivals in Cajas. Though they are currently
restricted to commensal activities, there is the danger that, as has occurred

29 specimens reported as Caenolestesfuhginosus by Barkley and Whittaker (1984) from Peru have been re-
identified as C. caniventer (see Pacheco et al., 1995).



elsewhere (e.g., Fox and Pople, 1984; Goodyear, 1992), these alien rodents may
begin to expand their range at the expense of the native species.
Human activities.-Unlike areas of lowland Ecuador (see Suarez et al.,
1995), small mammals in Cajas are not hunted for food. A number of indirect
impacts, however, can be observed and anticipated, though not yet quantified.
Agricultural activity occurs within the park. The impacts of ranching are of
particular concern. In the paramo of Cajas, as elsewhere in Ecuador (Knapp,
1991), the graslands are grazed by cows and burning to provide fresh pasture is a
regular occurance (Huber, 1979; Ramsay and Oxley, 1996). For example, at
Ventanas, a heavily grazed locality, no Phyllotis were caught. This was despite an
abundance of rock outcrops in the close-cropped pajonal, of the kind normally
favoured by this herbivorous rodent.
Burning results in loss of cover and food for rodents (Neal, 1970; Yates and
Lee, 1997) and has been shown elsewhere to cause a temporary depression of
rodent numbers and species diversity (e.g., Delany, 1986). Unless burning is so
frequent as to cause habitat degradation, the impacts of such fires are usually
temporary (Swanepoel, 1980; Kern, 1981; Rowe-Rowe and Lowry, 1982; Happold
and Happold, 1989). Such perturbations are a common feature of high elevation
habitats in the northern Andes (Armstrong and Macey, 1979; Lojtnant and Molau,
1982; LandAzuri and Jij6n, 1988; Lopez-Arevalo et al., 1993; Homewood, 1996),
where humans have practiced fire-management of the region's grasslands for
millennia (see Chapman, 1926; Knapp, 1991; Balslev and Luteyn, 1992;
Homewood, 1996). Post-fire vegetation regeneration patterns reported by Ramsay
and Oxley (1996) indicate that if the practices are operated at sustainable levels the
effects on the Plateau's small mammal fauna may continue to be minimal at the
meta-population level.
Quenoa forests are fragile ecosystems (Fjeldsa and Kessler, 1996). and
ranching activities may have severely impacted the quenoa forests of Cajas (Huber,
1979). Cows use them for shelter and in the process both trample the groundcover
and destroy the natural windbreaks of Diphlostephium and Gynoxis. This causes
desiccation and allows ingress of grassland plants. Both retard regeneration by
Polylepis seedlings (see Smith, 1977) and together these three effects cause
progressive loss of cover by this already localized habitat.30 Five of Cajas' small
mammal species appear to be restricted, or nearly so, to quenoa forest. Two (M.
altissimus and T. pyrrhonotus) appear to favor degraded and/or disturbed forest,
while C. montivaga, T. cinnameus, and T. gracilis are conspicuously absent from
heavily disturbed forests, and will even be absent from those parts of an as-yet still
closed-canopy quenoa forest that are most heavily visited by cows or humans. The
third threat comes from tourism. Cajas is officially designated as a recreational
area and the use of Cajas for such purposes has risen in the last few years (see Ashe

30 The extent of loss of Polylepis cover in Cajas can be guaged from the fact there are several place names
that specifically mention this forest type (e.g., "Quinoascocha" 'polylepis lake'), that now no longer have
any such forest in their vicinity.




Figure 10. Fif Robinson with a mountain coati, Nasua olivacea; shot by hunters at Yakatuviana (2400m).
Photograph, Adrian Barnett.


and Leschen, 1995; Barnett, 1997a). The effects of large numbers of humans to
Cajas has not been quantified but such impacts are rarely positive for mammals
(see Yalden, 1990; Gander and Ingold, 1997) or the vegetation cover on which
they ultimately depend (e.g., Bayfield, 1980). The effects of trampling can be
especially severe on moorland-like vegetation types, including paramo (see Brown
et al., 1978; Bayfield and Brooks, 1979). The use of Polylepis for firewood at
favored camp sites can have a very severe local impact (i.e., Lake Luspa) (see
Huber, 1979). Littering and fecal pollution are collateral impacts of unexplored
severity. Hunting of larger mammals (e.g., deer, Odocoileus and Pudu, and small
carnivores like the mountain coati Nasua olivacea, see Fig. 10) and trapping for
rabbits (Sylvilagus brasiliensis) have been observed during the study on several
occasions. The lack of comparative data makes it difficult to assess the severity of
these impacts, but it is suggested that a monitoring scheme be initiated.

Other Mammal Species

Other mammals species recorded on the Cajas Plateau were: Andean
Spectacled Bear (Tremarctos ornatus; piramo), Mountain Coati (Nasua olivacea;
cloud forest), Weasel (Mustela frenataa?] cloud forest), Hog-nosed Skunk
(Conepatus chinga; cloud forest), Mountain Lion (Puma concolor; pAramo and
cloud forest), Brocket Deer (Mazama rufina; cloud forest), White-tailed Deer
(Odocoileus virginianus; cloud forest edges), Porcupine (Coendou bicolor; cloud
forest), Mountain Guinea-pig (Cavia porcellus, pAramo), and Rabbit (Sylvilagus
brasiliensis, pAramo and grassland in cloud forest zone). There were unconfirmed
reports of Jaguarundi (Felis yagouroundi) and Northern Pudu (Pudu
mephistopheles) at Mazin. Long-term residents reported the Andean fox
(Dusicyon culpaeus) as formerly plentiful in the area, but now absent due to
hunting pressure.
The following bats were recorded in the Rio Mazin valley by Fif Robinson
(see Robinson and Hancock, 1986): Anoura geoffroyi, Myotis aff. oxyotis, Sturnira
erythromos, Histiotus montanus colombiae, and Desmodus rotundus. Myotis keaysi
keaysi was recorded nearby by LeVal (1973).


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Cryptotis montivaga Anthony, 19211 *
Caenolestes caniventer Anthony, 1921 *
Caenolestes filginosus (Tomes 1863) # centrahs, obscuru
Caenolestes tatei Anthony, 19232*
Akodon mollis Thomas 1894 altorum, fiilvescens.
Akodon orophilus Osgood, 19133*
Anotomys leader Thomas, 1906 #
Chibchanomys orcesi Jenkins and Barnett, 1997 *
Ichthyomys hydrobates Winge 1891 # nicefori, soderstromi.
Ichthyomys stolzmani Thomas 1893 # orientahs.
Ichthyomys tweedii Anthony., 1921 # courinus.
Neusticomys monticolus Anthony, 1921 #
Microryzomys altissimus (Osgood, 1933) chotanus, hylaeus,
Oryzomys altissimus.
Microryzomys minutus (Tomes 1860) dryas, fulvirostris, humilor,
Oryzomys minutus
Oligorzomys destructor (Tschudi 1844) maranonicus, melanostoma, spodiurus, stolzmani.
Oryzomys albigularis (Tomes 1860) caracololus, child, maculiventer, meridensis,
oconnelli, pectoralis, perrensis.
Oryzomys auriventer Thomas 1890 # mnmbosus
Phyllotis andium Thomas, 1912 fruticolus, meliamus, stenops, tamborum.
Phyllotis haggard Thomas, 1908 elegantulus, fiscus.
Rhipidomys latimanus (Tomes 1860) # colaensis, micotts, mollissimus, pictor.
Sigmodon inopinatus Anthony, 1924 hispidus.
Thomasomys aureus (Tomes 1860) altorum, nicefori, popayanus, creator, princeps.
Thomasomys baeops (Thomas 1899) *
Thomasomys cinnameus Anthony, 1924n gracilis, hudsonm.
Thomasomys gracilis Thomas, 1917 cinnameus, hudsom.
Thomasomys paramorum Thomas 1898 *
Thomasomys pyrrhonotus Thomas 1886 auricularis.
Thomasomys rhoadsi Stone, 1914 # fiumeus
Thomasomys silvestris Anthony, 1914 #
Aepomys lugens (Thomas 1896) # ottleyi, vulcani

According to Vivar et al. (1997), to be etymologically correct, the specific epiphet should be 'monivagus' so as to agree with the gender of
the generic name, Cryptotis.
Considered by Wilson and Reeder (1992) as a synonym of Caenolestesfuliginosus
Formerly considered a sub-species ofA. mollis, A. orophilus is now considered a valid species with two sub-species A. o. orientalis and A.
3. orophilus (see Patlon and Smith, 1992).
Thomasomys cinnammus was listed as a synonym of T. gracilis by Wilson and Reeder (1992), but Robert Voss (pers. comm.) considers it
to be a valid species His opinion is followed here.

Note: When referring to earlier literature, it may be useful to first check the taxonomic histories compiled by G.H H. Tate. These include
Phyllotis (1932b), Oryzomys (Oryzomys) (1932c). Oryzomys (Ohgoryzomys, and others) (1932d), Rhipidomys and Thomasomys (1932e)
andAkodon (19320



Collection numbers for skins collected by the author in Cajas and deposited in
the collections of the Natural History Museum, London1

/number of individuals

Cryptotis montivaga Anthony, 1921 N = 9 84.385, 84.386, 84.387, 84.388, 84.389, 87.917,
87.918, 87.919, no number.
Caenolestes camventer Anthony, 1921 N=1 84.383
Caenolestes tatei Anthony, 1923 N=1 84.384
Akodon mollis Thomas 1894 N= 108 82.752, 82.755, 82.756, 82.759, 82.760; 82.761,
82.762, 82.763, 82.764, 82.765, 82.766, 82.767,
82.768, 82.769, 82.770; 82.771 82.772, 82.773,
82.774, 82.775, 82.776, 82.777, 82.778, 82.779,
82.780; 82.781, 82.782, 82.783, 82.784, 82.785,
82.786, 82.787, 82.788, 82.789, 82.790; 82.791,
82.792, 82.793, 82.794, 82.795, 82.796, 82.797,
82.798, 82.799, 82.800; 82.801, 82.802, 82.803,
82.804, 82.805, 82.806 82.807, 82.808, 82.809,
82.810; no number, 84.224, 84.225, 84.226,
84.232, 84.233, 84.234, 84.235, 84.236, 84.237,
84.238, 84.239, 84.240; 84.241, 84.242, 84.243,
84.244, 84.245, 84.246, 84.247, 84.248,
84.289,.84.250; 84.251, 84.252, 84.253, 84.254,
84.255, 84.256, 84.257, 84.258, 84.259, 84.260;
84.261, 84.262, 84.263, 84.264, 84.265, 84.266,
84.267, 84.268, 84.269, 84.270; 84.271, 84.272,
84.273, 84.274, 84.275, 84.276, 84.277, 84.278,
84.279, 84.280; 84.281, 84.282, 84.283, 84.284,
84.285, 84.286, 84.287, 84.288, 84.289, 84.294,
84.298, 85.882,
Akodon orophilus Osgood, 1913 N = 9 82.750; 82.753, 84.290; 84.291, 84.292, 84.293,
84.295, 84.296, 84.297,
Chibchanomys orcesi Jenkins and Barnett, 1997 N=3 82.815, 82.816, 84.349.
Oligorzomys destructor (Tschudi 1844) N=5 82.847 84.299 84.300 84.302 84.303
Oryzomys albigularis (Tomes 1860) N=7 82.725, 82.726, 84.304, 84.305, 84.306, 84.307,
Microryzomys altissimus (Osgood, 1933) N=27 82.829, 82. 732, 82. 733, 82. 734, 82. 735, 82.
736, 82. 739, 82.740; 82.741, 82.742, 82. 743,
82.744, 82.745, 82.746, 82.747, 82.748, 82.749,
84.301, 84.309, 84.310 (twice), 84.312, 84.313,
84.314, 84.316, 84.317, 84.318,
Microryzomys minutus (Tomes 1860) N=2 82.731, 84.311
Phyllotis andium Thomas, 1912 N=1 82.813
Phyllotis haggardi Thomas, 1908 N=6 82.811, 82.812, 84.344, 84.345, 82.346, 82.347
Sigmodon inopinatus Anthony, 1924 N=4 82.814, 84.348, 85.884, 85.885
Thomasomys aureus (Tomes 1860) N=5 84.322, 84.323, 84.324, 82.325, no number
Thomasomys baeops (Thomas 1899) N=5 84.327, 84.330; 84.336, 84.337, 84.412.
Thomasomys cinnameus Anthony, 1924 N=6 84.315, 84.339, 84.340; 84.341, 84.342, 84.343,


Thomasomys gracilis Thomas, 1917 N=21 82.817, 82.818, 82.819, 82.820; 82.821, 82.822,
82.823, 82.824, 82.825, 82.826, 82.827, 82.828,
82.829, 82.830; 82.831, 82.832, 82.833, 82.834,
82.835, 82.837, 82.838.
Thomasomysparamorum Thomas 1898 N=13 82.839, 82.840; 82.841, 82.844, 82.845, 84.328,
84.329, 84.331, 84.332, 84.333, 84.334, 84.335,
Thomasomys pyrrhonotus Thomas 1886 82.319, 82.320; 82.321, 82.842, 82.843

1 The numbers in Appendix 2 refer only to skins. For some specimens only the skulls were preserved or
accessed into the NHM collection. This accounts for the occasional mis-match between the numbers reported
here and the species total numbers given in the text.

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