“Afiaafiu Mal-$2; “GWEN
35884.3 {333$ mam-4.314537%

20"! «‘9‘:

66%. $3.833

3": .n AmE'IVK (13-; {.1232 'Rc._c1r,nfi 3 :3; ‘
‘ .'..,. ’ '. A ‘3" '1‘.‘ g.- ,
“Luv-K. slaw..- J. I .c: and 9V5;JU 3. I Lil I”.

""'T .. Fag—”1’. airy! .‘us flit!

.v , - 1 st'v .- ‘) -.. ,
-. -. t :-;_ v '-n L_, I ion-v ., .
) ;}‘.— _ ;‘-" . 3.... . .I.‘ .
.....J..-\...... (5...... V.-..-—.t~‘-. -

fl_=1r\_f;5n' ~_ ‘P’I-“pu- v
3- . ' g;u.u.‘~:. 3.32
~ u

~U0:WI-‘; J ...-.-'.‘.
r: n-gn
lain
“if c

 
  
 

.~ I. 5‘. I!

  
   
   

LIBRA R Y
Michigan Stab

    
 

This is to certify that the
thesis entitled
MAMMALIAN-SIPHONAPTERAN ASSOCIATIONS
IN SOUPHHESTERN COLOMBIA
presented by

Eustorgio Héndez

has been accepted towards fulfillment 1
of the requirements for 3
l

Doctoral degree in Philosophy

 

l
DQM .
Dr. Rollin H. Baker

Major professor

Date Mag 28, 1976

0-7639

ABSTRACT

W‘MMALIAN—SIPHONAPI’ERAN ASSOCIATIONS
IN SOUI'HWESTER‘I COLOMBIA

BY

Eustcrgio Méndez

A synopsis is presented of the flea fauna and their mammalian hosts

in sout‘rmestern Colombia, with particular reference to the Departamento

del Valle. The information existing on the Siphonaptena of Colombia is

reviewed. Four new species of the genus Polygenis Jordan are described.

In addition, descriptions are given of the male of Ctenidicscmus traubi

Johnson, and the female of Sphinctopsylla diomedes Johnson. The former

species was previously known from the female, and the latter species was
originally described from the male. The following taxa are reported for

the Republic of Cclonbia for the first time: Plocopsylla phyllisae

Johnson, Leptopsylla segis (Sdmonherr), Dasypsgllus gallinulae

 

Eginnatts (Baker), Tetrapsyllus ccmis Jordan, Polzgenis pradoi (Wagner),
2. thtmmani Johnson, 2. kligesi samuelis (Jordan and Rothschild), and
Pulex simulans (Baker). The taxon Rhyndwpsyllus megastiggnata Traub and

Ganmons, 1958, is regarded here as a synonym of Rhyncli@sylltzs pulex
Haller, 1895.

Records contained herein bring the total number of known Colcrrbian

Siphonaptera to at least forty-four species and subspecies. Of this

number more than two-thirds are reported from the southwestern part of
the comm.

Eustcrgio Mendez

A key to the fleas of southwestern Colombia has been prepared,

including illustrations of the majority of the taxa mentioned.

A succinct account is given of the history and zoogeography of
the Hmmalian fauna of southwestern Colombia, as well as comments on

host—parasite relationships . Geographic aspects , such as topography,

geology, soil, climate and vegetation , concerned with the pertinent biota

are discussed.

MAMMALIAN-SIPHONAPI’ERAN ASSOCIATIONS
IN SOUI'HWBSTERN COLOMBIA

u/ fi
(\V

H‘- \
u
\r

Eustorgio Mendez

A DISSERTATION

Smeitted to
Michigan State university
in partial fUlfillment of'the.requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Zoology
19 76

ACMOWIEDQ‘IENIS

The guidance, encouragement and valuable suggestions by
Dr. Rollin H. Baker, Director of the The Museum,Mid1igan State University,
are deeply appreciated. I wish to thank Drs. Harold D. Newson, James W.
Butcher, and Marvin M. Hensley of Michigan State University and Drs. David
C. Baerg and Richard N. Rossan of Gorgas Memorial Laboratory for their
assistance and critical review of the manuscript. Advice and support were
also given by Drs. Martin D. Young and Pedro Ganndo V., former and present
Director, respectively, of Gorgas Memorial Laboratory.

This work could not have been undertaken without the cooperation of
Dr. Harold Trapidc, who submitted the basic material and continued to
support the project following the termination of his assignment with the
Rockefeller Fomdation in Colombia.

I wish to express my sincere thanks to Drs. Alvaro Duefias and Maurice
Thomas for authorization to use information pertaining to mammal specimens
deposited with the Departamento de Microbiologia, Universidad del Valle .
Dr. Thomas has been kind enough to collaborate in other aSpects of this
work and has donated material .

1». Alberto D'Alessandro and other authorities, Drs. Pablo Barreto,
Stephen Ayala, and J osé I . Borrero , of the International Center for Medical
Research and the Universidad del Valle, as well as Dr. Vernon 13. Thatcher,
formerly with these institutions , have contributed in several aspects to

my investigations .

Dr. Eric C. Wells, head of the Animal Health Program of the Centro
Internacional de Agriculture Tropical at Palmira, and David E. Evans, of
the same mit, have provided flea specimens and needed information.

I am indebted to Dr. Hugh Tyndale-Biscoe of the Australian National
University, Canberra, for the loan of Colombian flea material obtained in
connection with his research on South American marsupials.

Dr. F. G. A. M. Smit of the British Museum Department of Entomology
kindly examfired specimens of the new taxa of Pclygenis and confirmed their
status as undescribed species.

Assistance in field activities at Departamentc del Valle was given
by Messrs Lucio Velasquez and Jairo Ardila, who are with International
Center for Medical Researdi.

Drs. Oscar A. Gonzalez C. , Bagardc Materén Narvaez and Jaime Vargas
of the Corporacién Auténcma Regional del Cauca helped make possible our
trip to Alto Anchicaya and afforded facilities for which I am indebted.
Thanks are expressed to Mr. Harold Graham for the hospitality offered
us during the collecting trip to Lago Calima.

Other persons who have assisted in this work are too numerom to
be listed individually. To each of them, I express my gratitude for their

contribution toward the completion of this work.

iii

TABLE OF CONTENTS

Eage-
LISTOFTABIES v
LISTOFFIQJREMW. .............. ........... vi
mrmonmom ........... 1
GBOGRAPHICDBSCRIP‘I‘IONOFTHB

DEPARmm'onmvmrs 6

TOPOGRAPHY ...... . .............. 5

sons ........... . ..... 12

VBGBI‘ATIONAL PORMATIONS.............. ........ . ......... . l8
PEES‘I‘ORICANDZOOGEDGRAPHICSWRYOF‘I‘IE

WOF SOUI'l-IWESTERNCDLOMBIA 26
Accomrsornmsorsormmmcomm.... ............... 39
ANALYSIS 0FI-DST-PARASITEREIATIONSI-IIPS...................... 39
IGY'I'OTI-IEHEASOFSOUITMSI'ERNCOLOMBIA.. .............. ....1ou
BIBLIOGRAPHY.... ........ ...... ......151

iv

Number

 

LIST OF TABLES

List of mammal hosts and their fleas.

Geographic affinities of the genera of
Southwestern Colombian fleas .

Geographic affinities of the species of
Southwestern Colombian fleas .

101

102

 

10

12
13
1H
15

LIST OF FIGURES

Map of Colombia

Rainshadow in transect of Departamento del
Valle

Soil map of the Departamento del Valle

Vegetatimal Formations of Departamento del
Valle

Dispersal Centers related to the southwest
Colombian fauna

General appearance of female of Ctenocephalides

 

fie—Jig (Bouche)
Structures of apex of aedeagus of Scolopsyllus
colcmbianus Mendez
Neotyphloceras rosenbergi (Rothschild)
Adoratopsylla intermedia copha Jordan

Ctenidiosomus traubi Johnson

 

Cleopsylla nonticola Smit
Sphinctopsylla dicmedes J ohnscn
Plocopsylla phyllisae Smit

Plocopsylla thor Johnson

Sternopsylla distincta speciosa Johnson

vi

38

112

113

1114
115
116
1.17
118
119
120
121

LIST OF FIGURES (Cont'd)

"Ii—.3321;

16
17
1e
19
20
21
22
23
2a
25
25
27
28
29
30
31
32
33
31+
35
36
37

I._eptopsylla segnis (Schcnherr)
Dasypsyllus gallinulae minnatus (Baker)

 

Pleochaetis smiti J ohnscn

 

Tetrapsyllus comis Jordan

 

Polygenis bohlsi bohlsi (Wager)

 

Polygenis caucensis, n. sp.
Polygenis caucensis , n. sp.
Polygenis delpontei , n . sp .

Polygenis deficntei , n. .sp.

 

Polygenis delpontei , n . sp .

 

Polygenis dunni (Jordan 8 Rothschild)

 

Polygenis hopkinsi , n. sp.

 

Polygenis h_opkinsi , n . sp .

 

Polygenis klagesi (Rothschild)
Polygenis pradoi (Wagner)
Polygeiis roberti beebei (I. Fox)

Polygenis thurmani J ohnscn

 

Polygenis trapidci , n . sp .

Polygenis trapidoi , n. sp .

Scolopsyllus colonbianus Méndez

Scolopsyllus colcmbianus Mendez

Rl'opalopsyllus australis M Jordan
8 Rothschild

122
123
12”
125
126
127
128
129
130
131
132
133
13”.
135
136
137
138
139
190
1H1
192

1M3

LIST OF FIGURES (Cont'd)

Number

38

39
no
ul
1.2
I43

nu

Rhopalgpsyllus cacicus saevus Jordan 8
Rothschild

Rhopalopsyllus lugubris Jordan 8 Rothschild

Xenopsylla cheopis (Rothsdiild)

 

Ctenocephalides felis (Bouche)
Pulex irritans Linnaeus
munchqpsyllus pulex Haller
Tunga penetrans (Linnaeus)

.Oi

11m

MS
1146
1147
1148
1149

150

INTRODUCTION

The Departamento del Valle, one of several political divisions of
the Republic of Colombia, comprises an area of about 20,Lt30 km2 and is
located in the southwest sector of that country. It contains contrasting
ecological situations , including the very wet rainfcrest of the Pacific
lowlands , the western and eastern slepes of the Cordillera Occidental,
the basin of the Rio Cauca lying in the rainshadow of Cordillera
Occidental at 1000 meters, and the ascending western slope of the high
Andean Cordillera Central. This territory is bordered on the north by
the departamentos del Chocc and Risaralda, on the south by the
Departamento del Cauca, on the east by the departamentos del Tolima and
Quindio, and on the west by the Pacific Ocean.

The Departamento del Valle is known to possess a great variety of
plant and animal life, representing more than half of the total number
of species of naturals known to occur in, Colombia. Despite the [diversity
of the fauna and the importance thay many of these animals have as positive
or negative elements in the health of humans and the domesticated animals ,
there is a paucity of information about them. This need for basic research
in biology is indicative of the whole of Colombia as well as other areas
of South America.

During the past two decades, the Universidad del Valle, in
conj motion with the Rockefeller Foundation and the Tulane University
International Center for Medical Research, have contributed significantly

1

to the knavledge of arthropods and mammals associated with zoonoses in
Colombia (Cali Virus Laboratory, 1965, 1968, 1970). Through the study of
ectoparasites collected by these research units, an interest developed for
the investigation of fleas and their mammalian hosts in the Departamento
del Valle and other regions of Colombia.

The importance of fleas in the epidemiology of plague, tularemia,
munine typhus and other diseases is well known, thus primary consideration
of these medically important arthropods is warranted. In some instances ,
a lclowledge of the ectoparasitic fauna of an area may also provide
mderstanding of the relationships among animal host groups (Clay, 1951;
Hopkins, 1992; Patterson, 1957). Although plague, which remains a major
concern, has not yet been found in Colombia, it is present in the adjacent
comtries of Ecuador, Venezuela, Peru and Brazil. There is, however, a
high potencial that the disease can appear in Colombia, since ideal climatic
factors occur in many areas of the country. In addition, there is the
presence of both wild rodent species and those commensal with man, that
have been implicated elsewhere either as actual or potential reservoirs of
plague. Several species of fleas occurring in Colombia, such as
Xenopsylla cheopis, the classical vector of plague, as well as _P_u_l;e)_g
irritans and certain members of Polygenis , have been implicated in the
transmission of plague in other countries (Chabaud, 1947; Macchiavello,
19%, 1951+, 1958; M011 and O'Leary, 191:5; Panamerican Health Organization,
1956; Pollitzer, 1951+).

Presently , the Siphoraptera fauna of Colombia is poorly known . In
a survey by the author prior to the begirming of this study it was found

tl'at Colombian material was not well represented in the collections of

several institutions , including the British Museum and the United States
National Museum of Natural History.

The literature contains few reports concerning Siphonaptera from
Colombia. Dunn (1929) and Patifo-Camargo (1990) briefly mention sore
common species. Fuller (19142) and Macchiavello (1998) gave several
Colombian records of fleas . Costa Lima and Hathaway (1916) present
information on several Colombian species. Gast Galvis (1950) in his
list of fleas from Colombia considered 19 species and subspecies.
Johnson (1954 and 1957) has most significantly contibuted to our
knowledge of the fleas of Colombia. In 1951;, she described a new species
of Pleoghaetis from that county. Her outstanding monograph "Pleas of

 

South America," published in 1957, contains several new distributional
records of Species for Colombia and, in addition, the description of six
new species. Mendez (1968) described a new genus and species of
Colombian flea, and more recently, Méndez and Hanssen (1975) reported

a new Colorbian taxon discovered in the Departamento del Meta. Tamsitt

and I. Fox (19 70) have presented records of Rhynchopsyllus pulex Haller.

 

This work is largely based on informatim obtained from collections
made in the Departamento del Valle and neighboring political divisions
of southwestern Colombia, such as Narifio, Cauca, Putumayo and Huila. In
view of the ecological affinities of these territories and the similarity
of their mammalian fauna, it is likely that the ectoparasitic fauna are
virtually the same.

The present account contains descriptions of four new species of
Polygenis Jordan. The female of Sphinctopsylla diomedes Johnson, mich

 

had been known only from the male, as well as the male of Ctenidiosomus

 

taubi Johnson, are also described. A key to the species of fleas known
or presumptively existing in the southwestern portion of Colombia, and

new records for the comtry, are also presented. Rhmchcpsyllus

 

megastigmata Traub and Gammons is considered here to be a synonym of

 

_R_. £1.93. Haller. Conventional figures for the majority of the species
concerned are included.

The nucleus of the material was from collections by Dr. Harold
Trapido while engaged in virus research sponsored by the Rockefeller
Foundation. Additional specimens were either personally collected or

obtained from the Universidad del Valle and other sources.

 

PAIAIA

PACIFIC
OCEAN

 

- .\ I,
\ , nuns .-
-.\ ° ‘3‘. . \'\ ' \ ”\0 ‘
0.‘.. \o-\fl «can .\'v~. . , J5 .
a. I

ICUADOI

PO

- / I / \ / / .,/ \_ _, V i
/::// ' “y / . -~ 0
(’7 . I . .r.m-A. .
%/ /./.~-‘_//-i one I {J’— I
" ' ... ._ I I
“on 0-13 -jv.|.\‘\ J I ’5' “AK “Mums

. J \

,\ um“ / \. l ‘
> < /' \'\ l ‘7‘” .~ ls," \
kt, ) K /' \ \ / \."I — .....

4" r. ! ’

ATLANTIC OCEAN \ ‘9

. .3 3:. fl} I,» . .f‘ v: uezueu
) l.” - - . '
\ (r

t
\ t m . ~.-I.om..3 "a" .
) I I“ I A Isa-nu...)
.- / l 1' \. V" j =
\ V" / l. ' ' . .
\r.

W \_ - I' / I
H / .

/_,. «I \, I \. (\. /.’

i P'.’ W \ ”race /'

GA ) $M_ '/'
. ( n. P- )g7\ f"./| "nun
/ . .~' .I. , )\ . . J, _ .

 

    

IIAIIL

 

 

 

 

 

REPUBLIC OF COLOMBIA

Figure 1.

GEOGRAPHIC DESCRIPTION OF DEPARTAMENTO DEL VALLE

'IOPOGRAPHY:

 

Irasmuch as most of the siphonapteran material used in this study
pertains to the Departamento del Valle , the following geographical
discussion is limited to this territory. No other political division of
Colombia displays such a diversity of geographic features and,
consequently , ecological conditions . The topographic information given
below is essentially derived from Espinal (1968), Sénchez (1965) and
Sater (1950 ) .

Generally, the physiographic and faunistic elements found in this
area are characteristic of the southwestern zone of Colombia, which
consists of the Cauca Valley territory, the surrounding mountains, and
the Pacific coastline within the limits of the departamerrtos del Valle,
Cauca, Huila, Narifio, and Putumayo.

The topographic scene is dominated by the Western and Cental
ranges of the Andean mountains system. These mountains contain some of
the higher peaks of Colombia. The Western Cordillera extends
north-northeastward parallel to Cental Cordillera and is almost parallel
to the Pacific coast. It is the lowest of the three ranges forming the
Andean system. The Cental Cordillera in the eastern sector of B1 Valle
contains hign peaks such as the mountains of Huila and Barragén, which

exceed 3000 meters .

' Several rivers and numerous steals form a Pacific watershed which
originates in the mountains and empty into the Pacific Ocean. The more
important rivers are the Anohicayé, Dagua, Naya and Cauca. This last:
river is the largest and courses througln the Departamento from south to
north.

Mangrove swamps flooded by higln tides are found along the shoreline
of the Pacific coasts . Warm topical rain forests follow the swamps and
occupy extensive inland zones of dense vegetation, which is impenetable
in many areas. The rich plant life, water, and cover, offer optimal
conditions for an abundance of animal forms . 'Ihe pocket-like arid valley
of Dagua, horever, with its grassy and brushy cover, introduces a
contesting zone in this lowland territory.

A subtropical zone of humid temperate climate follows the lowland
cloud forests. This temperate zone is characteristic of the middle slopes
of the mountains, where vegetation is rich but not very lnouriant. Many
of the fennel elerents of these subtropical forests are similar to those
found in the lowlands , since many species evolved from ancestors which
moved to the upper zones . Apparently the climate and other geographical
conditions have been limiting factors in the establishment of sore species
that have a more discriminating physiology.

The Cauca Valley, an agriculture area of the temperate zone, is a
mow isolated region consisting approdmately of L100 ,000 hectares lying
between the Western and Cental Cordillera. It is nearly 160 lciloreters
long and only 12 lciloreters wide, and is drained by the Cauca River.
Although mob of its territory is occupied by pasture, mainly Para
(Baum barbinode) and Guinea (2. madman) grasses, the Cauca Valley

‘-

2w..

~-

I

 

represents the most fertile and productive agricultural land of Colombia.
Lush montane cloud forests occupy extensive zones of moderate and
high elevations (from below 1000 to about 3000 meters) throughout most of
the Cental and Western Cordilleras . The uppermost reaches of the
mountains, from over 3000 meters, are largely unforested, consisting
primarily of extensive grass plains characteristic of paramos (such as
Las Hermosas, Chinclne, Miraflores and Barragén). These areas support more
selected types of plants and animals; indeed the paucity of animal life in
the paramos is directly correlated to the poor diversity of cover.
As in other areas of Colombia, the Departarento del Valle is
suffering from much deforestation, due primarily to agricultural

development, hydroelectric projects and cattle raising.

GEDIOGY:

By virtue of its rock composition, Colombia can be divided into
two geological regions , the Oriental plains , and the Andean geosynclinal
regions (Bu'rgl, 1961). Only the latter region is present in
southwestern Colombia, including the Departamento del Valle . The vast
Oriental plains are in east and southeastern Colombia.

The Andean geological region was apparently submerged during long
periods from the beginning of the Cretaceous , and accumulated large
deposits of marine, continental and volcanic sediment. The complicated
tectonic movements experienced by these lands were responsible for the
formation of the present Andean Mountains , which constitute the

"backbone" of Colombia.

The Andean geosynclinal region was consolidated before the Cambrian
period, and is divided into the following mountain systems: 1. The
Cental Cordillera, 2. The Western Cordillera and Coastal Cordillera
(Serrania de Baudé), and 3. The Eastern Cordillera. The lower and
Middle Magdalena Valley basins separate the Cental and Eastern Cordillera.
The Lower Valley contains non-marine Tertiary rocks , while , the Tertiary
reservoirs of the Middle Valley are non-marine . Western and Central
Cordilleras are separated by the valleys of the Upper Cauca and Upper
Patia rivers . Other mountains of Colombia, such as the Santa Marta
Mountains, Perijé Mountains and the Pacific Coast Range, have affinities
with the Andean system.

The coastal zone of the Andean Region is represented by the Bolivar
Geosynclire, a lowland area of Tertiary Marine formation, west of the
Andean mountains and extending from Southern Ecuador to the Gulf of
Urabé in northern Colombia. This strip of land has been interpreted as
a seaway which apparently permitted the moverent of terrestrial animals
during periods ranging from upper Cretaceous times to Recent (Nygren,
1930; Hershkovitz, 1966). It is of interest to note that no marine
formations from the Upper Cretaceous have been found in this zone .

The Eastern Cordillera displays abundant deposits of Cretaceous
and Tertiary rocks, with little or no recent volcanic elements. In
addition, Jurassic, Triassic and Paleozoic rocks are also found in these
mountains. Restern and Cental Cordilleras, as well as the Pacific Coast
Range, are primarily formed of igneous and metamorphic rocks , having
only subordinate sedimentary beds. In both Cordilleras, several high

volcanic peaks emist .

10

The Precambrian sedimentary history of Colombia is not known
(Jacobs, Biirgl and Conley, 1963). According to these authors, during a
great part of Cambrian and Ordovician time, the actual territory of the
Eastern and Cental Cordilleras , and at least the western aspect of the
Llanos and Putumayo-Caqueta lowlands , were occupied by seas . Later, the
area care under the influence of volcanic and diastophic activity, which
destoyed the Cambrian—Ordovician marine deposits. Rock shields ,
characteristic of the Andean system during the Cambrian , apparently were
greatly disturbed by erosion and their detitus filled up marine and
terrestial depressions of the region. The most important fossils from
the lower Paleozoic in Colombia are brachiOpods , trilobites and
graptolites . The territory west of Cental Cordillera shows no indication
of Paleozoic marine sediment.

Sore Middle Ordovician fossils have been found in Colombia. However,
Upper Ordovician and Silurian apparently are not represented by
fossiliferous layers in this county.

Fossils from the Devonian of Colombia are represented by brachiopods ,
bryozoa and trilobites , among other invertebrates . Many plant and animal
fossils, primarily marine forms, are known from the Carboniferots.

Evidence obtained from the fossils indicates that the lower
Carboniferous sea invaded the eastern part of the Andean Region, and was
covered by sediment thereafter. During the upper Carboniferous, the sea
gradually reteated and large semi-swampy forests moved to sections
previously occupied by water. Abundant Permian fossils have been found
in only a few areas of Colombia, primarily at Serrania de Perija. They

are represented by sponges , crinoids , brachi0pods , gasteropods ,

ll

cep‘ralopods, and other marine animals . The eastern portion of the
Central Cordillera contains Marine Upper Triassic rock, which has been
designated the Payande Formation, and represents a combination of
sandstone, limestone and shale. The Eastern and Central Cordilleras
contain lower Jurassic sedimentary rocks, principally of continental

It is evident that much igneous activity occurred in the Andean
Region during the Mesozoic. Also, it is apparent that the major part
of western Colombia was occupied primarily by seas during most of late
Jurassic and Cretaceous time. The fossil records from these periods
are scant and consist mainly of ammonites and other mollusca. The
Cretaceous Colombian and Peruvian faunas displayed a strong rela-
tionship with those of Southern Europe (Olsson, 1956). Considerable
tectonic movements , that occurred during Late Cretaceous and early
Tertiary, produced constant changes in the landscape of this region and
in the formation of marine and non marine deposits. The Tertiary
Continental deposits contain excellent vertebrate fauna , including
nurerous marmals. There is also evidence of considerable faulting and
folding during the late Paleocene .

Throughout the Eocene , geography of this area still was somewhat
different from the present. However, the principal elements of the
Andes and the Pacific Coast basin originated during this period. Weeks
(19%?) points out that during this period the Bolivar Geosyncline
introduced important changes in Western Colombia and Ecuador. Much
tectonic and volcanic activity took place during the Eocene and Oligocene

Gianna-n , 1961; Vuilleumier, 1971). Very few fossil vertebrates pertaining

12

to the Eocene are known from Colombia (Stirton, 1953). Layers
corresponding to the Eocene and Oligocene are richer in Eoramninifera,
molluscs and other marine fossils. The Cauca Valley is particularly rich
in Oligocene marine rocks, mainly consisting of algal, foramniniferal
limestone . This area also has coal—bearing Tertiary rocks , that probably
originated during the Miocene or before .

A large number of fossil vertebrates has been discovered in Miocene
deposits. The mammal fossil fauna of this period is very interesting and
contains some relics of families believed to have disappeared in earlier
times. Numerous findings of mammalian Pleistocene fossils have taken place
in Colombia (Stirton, 1963; Patterson and Pascual, 1968). Many of these
fossils represent species that became extinct at the end of the
Pleistocene. Jacobs, Burgl and Conley (l_o_c. _c__1_t .) consider that the most
recent stage of tectonic movement and volcanic activity was initiated in
the early Miocene and continued into Recent time , particularly affecting
tine Central Cordillera and the southernmost part of the Western

Cordillera. Some of the volcanoes existing today are still active.

SOILS :

The following considerations of the soil distribution of the
Departanento del Valle are based primarily on two major sources: the
FAG/UNESCO Soil Map of South America (1961), and the account by Beek and
Bramao (1968). In preparing this outline, an attempt has been made to
present it in a condensed form.

A great deal of work has been done in South America to determine

the distribution of major soils as important patterns in the development

13

of agricultural zones and the exploitation of minerals and other natural
resources significant to the general progress of the continent. However,
despite the knowledge that has been accumulated during many years , the
study of the structure and composition, as well as the distribution of
South American soils is far from being complete.

The nature of the soils of the Departanento del Valle is linked to
patterns of ecological factors that govern the life zones , such as climate,
geonorphology, topography, and vegetation. Departanento del Valle , and
other areas of southwestern Colombia, from which material used in the
present study has been collected, consist of Lowlands and Andes, two of
the major structural elements of soil distribution that have been
established for South America. Uplands, the other elements considered in
this segregation, are found in the eastern part of South America. The
criteria assured for the establishment of these general categories are
based on a complex association of factors such as geography, climate,
vegetation, physiography, etc.

The Pacific Coastal Lowlands form a high portion of the western
side of the Departamento and are primarily characterized by contiguous
areas of tropical evergreen and deciduous forests. The alluvial soils
predominating in this territory are stratified with little organic matter
and little profile development. The Pacific littoral is particularly
culminated by dense mangrove forests, interspersed with swarp forests. This
zone contains Quaternary marirne and fluvio-marine deposits with alluvial
plains and terraces , estuarine and delta alluvial deposits , and local

areas of coastal sand dunes.

11$

The Northern Andes is the other soil region found in the Departamento
del Valle. The soil composition is more diverse and the profile of the
area interrupted by the western and central ranges of the Andean mountains .
These are separated by the Cauca Valley, a strip of land which also
includes areas of the Departamento de Cauca, Narifno , Caldas and Antioquia.

In the western sector of the Northern Andes region of the Departa—
mento del Valle, volcanic rocks are the most important elements of the
soil structure. This condition reflects the volcanic origin of these
mountains . Most soils in areas of moderate elevation in the northern
Andes are Laterosols , derived from volcanic material. Laterosols are dark
colored surface soil with lighter subsoil, and, in addition to being
slightly acid, contain a high anount of organic matter. Because of their
low fertility level, they have limited use for farming. Extensive areas
of the northern Andes are represented by Andosols. Such soils consist
primarily of volcanic ash with dark surfaces , combined with organic matter
and minerals such as nitrogen, phosphorus , calcium and potassium. Those
that are not too acid are relatively fertile and excellent for growing
different crops . Sectors of the Departarento del Valle , dominated by
Andosols , have excellent pasture lands and areas devoted to agriculture .

Adjacent to Andosols are found the dark paramo soils, which may be
derived from heavy clays , perhaps of glacial origin. They consist of some
volcanic ash and are characterized by a high degree of acidity and paucity
of nutrients. The paramos lands have high humidity, low temperature, the

vegetation is poorly diversified and consist of pastures and forests of
secondary gowth.

15

SOIL MAP OF THE DEPARTAMENTO DEL VALLE

 

  
    
   
   
   
  
 

ANDOSOLS

ALLUVIAL SOILS

FOREST LATOSOLS

P RAMO SOILS

REOOISH BROWN LATERITIC SOILS

 

 
  

  
 

m,

L
3.31, '
t 1’

       

f a

 

 

 

 

Figure 2.

16

The northern part of the Departamento del Valle is characterized by
Reddish Brown Lateritics. These soils contain dark, reddish brown,
granular clay surface soil, with yellowish-brown , clay subsoil. Aluminum
silicate, its principal mineral corpound, is mixed with iron and other
ingedients of inorganic and organic origin. They occur below 2000 meters
and on which are found primary and secondary forests in addition to

pastures, coffee plantations and other cultivated areas .

CLD’IATES :

The complexity of its geography contributes to the variety of
climates existing in the Departamento del Valle, since these elements
are intimately associated. It is interesting to note that all four of
the climatic regions outlined by Eidt (1969) for South America, are
represented in the Departamento del Valle, namely, tropical rain,
terperate, arid and tundra.

Tropical rain climates are confined to the Pacific territory,
from sea level to the base of the mountains reaching an elevation of
close to 1,000 meters. These areas are hot and humid, with temperature
ranges from 24°C to 30°C. The annual rainfall is heavy, exceeding
760 on a year. It rains almost every day, thus this area does not have
a true dry season. The humid warm winds of the Pacific Ocean and the
Andean chain contribute to the heavy precipitation; Figure 3 , adapted
from Espinal (1968) , depict the rainshadow influence . According to
Espinal (Log. _ci'_t_) , within the Western and Central Cordillera there are
two rainy periods during the year: the first from April to June, and the

17

.m 95mm

 

 

 

 

 

 

 

(CWJJBCOO SFZMO >m44<> (030 (Gun—4.200 Icy—hm; Z<NOO 0.L_0(s
. learn/MN Mani. _
“Wu a:

iron)» 01.». mow/o 9...... .
as; _
w: _
_ . Z / _
_ ,,,l,,_~_a _ _ n
_ _ _ u _
_ _ n _ _
_ _ _ _ u
_ n _ _ _
_ _ _ _

_ _ _ _
_ _ _ _ _
_ _ .3. .
— g a; illusfl ilTluIONgog g3)! i—

 

333 an Omev<<h¢<me m0 hUmmZ<E 2. 3005.3 2.5.

 

'9“ .

J...

 

‘n-

‘-

‘-

.
on.

 

I
V

18

second from September to November. Between these wet seasons there is
notably decreased precipitation.

Tenperate climates are characteristic of the subtropical areas
extending from 1,000 m to 2,000 m, which have a temperature range of
18°C to 24°C. These areas are the first table lands found above the
lowlands, and are influenced by the higher masses of the Andes and by the
action of two kinds of winds , interpreted as mountain—valley breezes
and land-sea breezes . These winds effect the climate and precipitation
of the Cauca Valley system, the rivers, and other subtropical areas of
this territory.

The eastern part of the Departamento maintains a cooler climate
typical of areas above 2,000 meters, (in which various paramos are found).
The northern sector of the Central Cordillera contains an arid zone ,
subjected to strong winds . This zone contrasts with the cloudy and
rainy forests dominating the Andean landscape. Some of the higher
peaks of Central Cordillera are cool and damp paranos where the

terperature is below 12°C.

VEGETATIONAL FORMATIONS:

Most of the information presented below, including Fig. 11., is based
on the works by Espinal (1968), and Espinal and Montenegro (1963). These
studies were made according to the classification established by
Holdridge (19W), Which considers aspects of terperature and humidity in
the analysis of life zones.

19

The ecology of rodents and other mammals , as well as of the
ectoparasites affecting then, is highly dependent on biotic conditions ,
soils , climates and other factors . Vegetational formations are therefore
of primary importance in understanding habitat requirements and
distribution patterns of these animals. However, it must be pointed out
that such formations in time and space usually gade into one another
and therefore are not always permanent . In addition , many animals and
plants are able to tolerate a moderate range of climatic conditions .

A brief presentation of all of the vegetational zones outlined for
the Departamento del Valle follows .

Tropical Very Dry Forest. This Vegetational formation is distributed in
two areas. One is Loboguerrero, at the upper part of Rio Dagua in the
center of the Departarento. The other region represents a flat belt of
forest platform in the Central Cordillera, extending from Cali to near
San Francisco at the left shore of the Cauca River, reaching an
elevation of 1200 to lu00 m. The prevalent climate of this zone is dry
and the mean temperature is over 2l+°C. The mean rainfall during the year
ranges from 600 to 1000 mm. Characteristic plants of this zone are figs

(Ficus spp.), Vaclnelia farnesiana, spurge (Euphorbia caracasana) ,

 

Pithcellobiun dulce, Desmanthus virgatus , Achatocarpus nigricans , mesquite
(ProsoPis juliflora) , Jatropha gossypjifolia, Rngatera pterota, Ocimum

micranthum, Heliotropiun sp. , Lantana canescens, Citharexylum sp. ,

 

Portulaca pi losa and Talinun paniculatun.
Tropical Ey Forest. It is distributed along the Central Valley crossed
by the Cauca River and also as a strip of land surrounding the Tropical

Very Dry Forest of Loboguerrero. The mean tenperature of the area is over

. I. N . n!
.u a \nV \ ... w. fix an».

20

24°C and the rainfall fluctuates between 1000 to 2000 mm. The
characteristic vegetation of this zone grows in lands not over 1000 m
high and consists of cabuyas (aneraea sp.), m sp., T‘urnera
alnnifolia, Cephalocereta colonbianus and other plants .

Miml Moist Forest. This zone occurs in three areas below 1000 m

in the npperhalfofthe Departamento. They are the canyon of Rio
Garrapatas near El Cairo and Versalles, a strip near Rio Dagua and
PIObably an area in the mid portion of Rio Calima. The mean tenperature
is higher than 2u°c and the mean rainfall is from 2000 to ”000 mm. Some
Plants found in this type of forest are the balsa (Ochroma M),
mble (Tabebuia pentaphylla) , and hogplun (SEEM m).

Meal \Lgy Wet Forrest. This formation extends from the basin of Rio
Al'1<=hicay§asabelt t‘ratruns fronsoufintonorthalongtheWestern
CcI‘dnillera. The area is chrinated by a variety of plants such as ceiba
(Sign; m), balsa (Ochrora lagopus), black rubber (Castilla
Elastica), guaruros (m spp.), calabash (Crescentia c_ujete), cedars
(Came spp.), annato (g3 orellana), sandbox (Ego; crepitans , and
0there. These landsarenotabovelooo m. T'hetenperatmeexceeds

2lt°c and tlm is a mean rainfall of noon to 8000 mm per year.

M931- BaJn_’_ m. It is the largest of the vegetational formations
Ofthenepartamnmmactmdsfzmthevexywet lowlandcoastlinetoflne
nm‘gins of the Western Cordillera. Tfe dense mangrove forests consists
Miner-11y of red mangrove (Rhizophora brevistyla) in association with
black mangrove (Avicennia fl), white mangrove (Laguncularia racemsa),
and buttomood (Connocarpus). The inland forests, beyong the Pacific
littorial, contain a diversity of plants such as guaruno (M sp.),

21

Wild figs (figs), mammagia (Inophleum armatum), membrillo (Gustavia
M), Acacia melanoceras, sensitive-plant W) , cashew

(Anacardiun excelsun), balsa (Ochroma spp.), and many others. The

tropical rain forests are lowlands below 1000 m, having a hot

climate with a mean terperature above 21+°C and the mean annual rainfall
exceeds 8000 mm.

Sibtropical Dry Forest. It includes isolated areas on the eastern half

01’ the Departamento. The major area runs along the oriental baseline of
the Western Cordillera from Cali to San Francisco. One patch is found

in El Dovio's ravine and two other patches occur in the Central Cordillera.
Typical plants in this zone are figs (£i_c_u_s_ spp.), fiques (Fourcraea sp.),
mosCluero (933L911 sp.), big belly tree (Wigandia caracasana), indigo
(W sp. ) , cuipo (Cavanillesia platanifolia) , anong others. It
mgisters a mean temperature of 17°C to 21+°C and the mean annual rainfall
is between 500 to 1000 mm.

Subk'u‘opical Very Humid Forest. This formation encompasses very humid areas
Of the Western and Central Cordillera having an elevation between 1100 to
1900 m. The mean tenperature is from 17°C to 21+°C and the annual rainfall
fl~L1'Zl'tuates between 2000 to IJ.000 m. The vegetation of this zone is re—
pr'Q'Sented by guamo (1113?; spp.), guayacan (Tabebuia chrysantha),

‘OQmm lagopus), gnaruno (Cecropia), yaragua (Melinis minultiflora, pigs

 

 

fer}, (Pteridiun aguilinum), fox tail (Andropogon sp.), and others.
SubX‘lzopical Moist Forest. It is distributed as an extensive area
Suttounding the dry Rio Cauca valley and extending to the edges in the
w‘i‘i‘a'tern and Central cordilleras . Other patches of this zone are located

in areas surrounding E1 Dovio and in the upper portion of the rivers Dagua,

22

El Carmen and E1 T'reinta. The zone reaches an altitude between 1100 to
2000 m. The mean temperature ranges from 17° to 29°C, while the annual
rainfall is between 1000 to 2000 mm. Typical plants of this zone are the
macedero (T‘ricl'antera gigantea) , surr'umbo (Trema micrantha) , cordoncillo
(P_iper aduncun), balsa (Ochroma lagogne), iraca (Carludovica palmata),

 

trumpet (Bocconia frutescens), coralito (Hamelia patens), rubber (Egg
Sp.), chayote (Sechium edule) and many others.
S_Ub‘tr’opical Rain Forest. It is particularly represented by a strip of
Watered land of the Western Cordillera with elevations between 900 to
1900 m. The humidity in this vegetation formation is very high and the
temperature reaches from 17° to 2u°c. The annual rainfall exceeds uooo mm.
This zone combines sore virgin forests as well as agiculture lands . Some
Of its characteristic plants are platanillos (Heliconia), ferns
(Egdneniaceas), guarumo (Cecropia) , rubber (Castilla elastica), paco
(Egdesia macrophylla), pejibaye (Guilielma gasipals), and avocado
(m americana) .
3’73 Montae Dry Forest. This formation is limited to an area of high
elevations (from 2000 to 3000 m) of Barragan in the Central Cordillera.
The region possesses a dry climate with a mean temperature of 12° to
17°C, and an annual rainfall of 600 to 1000 m. The native vegetation
is poor and consists primarily of capers (9333.12 sp.) and mosqneros
(\Cl‘bton sp. ); cultivated species include maize (_Ze_a_ w) , onion
Cm“. um _<_:_ep_a_) , wheat (Triticun) , and potatoes (Solanum tuberosum) .
%_ Montane Moist Forest. This region consists of two areas in the
CeIntral Cordillera, where the altitude is from 1800 to 3000 m. It is a
Gold formation, the huridity is high and the mean temperature varies

 

 

 

23

from 12°C to 17°C. The annnal rainfall is over 1#000 mm. Sore of the
plants found in this territory are the following: carbonero (Bejiria sp. ) ,
chilco (Baccharis sp.), capers (_C_as_§_i_a sp.), and mortif'o (Miconia
albicans).

Lower Montane Very Wet Forest. It represents extensive subtropical lands
of the Western and Central Cordillera having elevations between 1800 to
3000 m. The cool climate characteristic of this formation registers a
mean temperature of 12° to 17°C. The annual precipitation is calculated
to be between 2000 to I+000 mm per year. These forests contain many
different plants such as guarumos (93m spp.), lichens (Cora pavonia,
Cetraria sp.), mosses (Polytricun sp.), horse tail (guisetum sp.),

 

 

amino (Weimnamia balbisiana), cascarillo (Ladenbergia), etc..
Liner Montae Rain Forest. This is a wet formation distributed as
Occasional areas in the Western and Cenual Corchillera. The elevation
I‘Emees from 1800 to 2900 m. and the mean temperature varies from 12° to
17°C. The annual rainfall registers over l#000 m. Typical plants of this
“me are the white gnaruro (Cecropia), capers (_C§_s_§_ig sp.), quassia
( sia sp.), cherry (Preziera sericea), cedrillo (Brunellia sp.), lulo
\S°1anun guitoense), horse tail (ransom bogotense), berries (3% sp.),
We (M sp.), and others.
Lantana vfl Wet Potent. This formation occupies a broad portion of the
WPicalparanodistributedfromSantaheiatoBanaganinflneOentral
cQ3!I'~clilla:a. It has an elevation above 3000 m and a mean temperature from
5" to 12°C. The anrmal rainfall fluctuates between 1000 to 2000 mm. The
1“ant-life is represented by. frailejones (Es letia, capers (_C_a_s_s_i_a_ sp.),
eSpender!) (Rama sp.), charcoal maker (ma sp.), morteno (Hespermeles

 

 

2”

sp.), grasses such as Calanagrotis and Festuca, etc.

Montae Rain Forest. These areas are very hurid, over 3000 m in altitude

 

which are localized in the Western and Central Cordillera. They are
paramos that have a mean temperature between 6° to 12°C and a rainfall
that is over 2000 m. This type of vegetation formation contains cultivated
territories as . well as undisturbed forests. Among the plants found in
these lands are frailejones (Espgetia sp.), blackberries (M sp.),
encenillo (Weinmannia sp.), Serecio sp. , Vaccinium sp. , cherry-tree

(Prune sp.) and Badnaris sp.

25

 

    
 
  
 
 
 
 
 
 
 

IIHIIHIII‘ TROPICAL VERv DRY FOREST
TROPICAL ORV FOREST
- TROPICAL MOIST FOREST

 

- TRORICAL RAm FOREST

- SUITIOPICAL DRY FOREST

m SUITRORICAL MOIST FOREST 0

- suaTROPICAL VERY WET FOREST

- suaTRORICAL RAIN FOREST

m LOWER MORTAR: DRY FOREST 0

- LowER MONTANE MOIST FOREST

- LOWER MONTANE VERY wET FOREST
1 LOWER MONTANE RAm FOREST

- MONTANE VERY wET FOREST

- MONTANE RAIN FOREST

VEGETATIONAL FORMATIONS °

‘Api

 
    
      
    
 
  

PACIFIC
OCEAN

  

2

 

 

 

DEPARTAMENTO DEL VALLE

Figure 1+.

HISTORICANDZOOGEDGRAPEHCSIWOFTHEMAPMALSOE
SOUIHWBSTERNCOLDMBIA

The ecology of mammals and their ectoparasites in Colombia and other
South American countries is of interest to investigators that are
concerned with a variety fo zooncses linked to finose animals. However,
little has been reported on their biology and distribution . Even the
systematics of finese vertebrates is not complete and many problers
exist with respect to the nomenclature and the status of a number of
forms.

An attempt to understand the origin and distribution of mammals
presently occurring in southwest Oolonbia mats with an array of
discussions by authorities on the origin of south American mammals. To
fit fine limited scope contemplated here, I have tried to be selective
in the interpretation of possible events that, in my Opinion, have a
logical foundation. Several references concerning fine evolution and
zoogeography of South American mammals are given in fine bibliography.
(See partictflarly Herahkovitz, 1962, 1966, 1969, 1972; Keast, 1968;
Loonis, 191'4; Patterson and Pascual, 1963, 1968; Savage, 197“;

Scott, 1937; Simpson, 19%, 19u3, 19%, 1950, 1969; SW, 1950).

Thefineoyofplatetectanicshasirrtroducedanewargment
for exploring the origin and distribution of Latin American fauna and
Flora (Raven and Axelmd, 1975; Valentine and mores, 197k). Acmrdilng

26

27

to finis fineory, fine movement of land masses or plates constituting the
earfin's crust produces cataclysms and volcanic action with subsequent
conplicated modifications of the elements involved. As applied to fine
southern continants, and particularly to South America, some intriguing
perspectives occur with regard to fine origin and affinities of the biota.
In View of fine geological facts , during fine Middle Cretaceous South
America was connected with Africa, and via Antarctica, wifin Madagascar,
India and Australia. Millions of yeas later it was gradually separated
from finose lamb . Apparently fine connection wifin Australia lasted longer.
It has bean speculated on finis basis that an exchange of fauna, at least
in part by island—hopping, took place betwnen Australia and South America
until fine Early Oligocene. 0n fine ofiner hand, a more effective connection
between South America and Africa existed nmtil fine end of fine Cretaceous.
It is apparent finat during fine Cretaceous fine separation bemoan South
MericaandAfrioavaswiderthanfine gap endstingbemeen South and
North America.

Probably during fine Late Bocane, fine interchange of tropical and
warm-tonperate animals between South America and Africa was greater
fianbetweenAfricaandNorfinAmerioa. Sonegroups ofanimals, suchas
fine river turtles, family Peloreducidae, which occur in Soufin America,
Madagascar and Africa, also seem to afford a logiml basis for finis
assunption. It is also believed that routes for the movement of
cool-temperate organisms between soufinern Soufin America and Australia
existed until nearly In million years ago. The argument appears to be
substantiated by certain elements of fine flora af angiosparms common
in Africa and South America.

28

Events of a different nature, primarily geological and climatic,
occurring in fine millions of years since the separation of fine southern
continents during fine Cretaceous , have interrupted fine faunal development
and fineir distribution in South America. This has introduced many gaps
finat obscure fine entire historical sequence .

Geological changes during fine Cenozoic, when fine mammals were in
fine process of development and evolution, are very important in attempu'ng
to understand sons of fine general history of fine mammals of fine Departa-
mento del Valle, and other parts of western Colombia.

The manlalian faunal structure of South America begun to evolve
wifin primitive forms recorded from fine Paleocene, such as marsupials,
members of the orders Condylarthra, Notungulata, Litopterna and other
mgIflAtes (Hershknovitz, 1968; Patterson and Pascual, 1968; Simpson,
1950).

Apparently during the beginning of fine Tertiary, fine Middle
American bridgepermittedmamnals tonnvetoandfromNorthand South
America. The many fossil discoveries from southern South America have
revealed fiat , early in the Paleocene , fine most important nuclei of
South American mammals were concentrated in finis portion of the confinent .
The earliest manual records from northern South America and Middle
America are represented only by marsupials and Condyla'ths .

During fine Eocene and Oligocene epochs, and presumably dnming
most of Cenozoic, Ncrfin America was isolated from South Anerica by a
seabarriersincefine connection representedbyfine Isfinnian linkwas
interrupted. Fossils of monkeys and cavionerph rodents lnave been found
in fine Oligocene of South America. However, fine evolution and dispersal

29

offinesemammalspriortofinis tnerenainamystery. Hershkovitz

(1968) and Simpson (lens, 1950) believe that these manuals invaded South
America probably as island-hoppers in fine aerly Oligocene. According to
Raven and Beelrod (_lgg. gilt), it is more probable finat primates and
cavionorph rodents reached South America from Africa during fine Oligocene,
whenfine distances separating NorfinAmerica andAfrica fronSouthAmerioa
were almost equal. .

Prom fine fossil history, it is evident that armadillos, anteater-s
and gound slofins inhabited South America since fine Tertiary and were
probably distributed in areas of mild climate, at leat close to our
present topical conditions. It is possible finat finey originated from a
Cretaceous stock. According to Simpson and ofiner authors, they gradually
moved into South America when fine Middle American land bridge was
re—established in fine late Pliocene. There is no indication that finese
mammals migrated between South and North America before finis time, with
fine probable exception of fine ancient armadillos. Fossil armadillos have
been discovered in early Tertiary heck of North America (Kiirten, 1969).
Hershkovitz (1972) discussed fine probability finat migation of terrestial
animals started during periods prior to fine land connection. Other
aufincr-s (Scott, 1937; Simpson, 191:0, among finen) maintain finat citing
fineperiodofisolationoffinecontnents, fineSoufinAmnericanroderrt
fauna was apparently represented only by caViotorphs . They also believe
thatfineNorthAmerican fauathencontairedmyomrphandsdaororph
rodents, but no cavioxorphs. According to Hershhwitz (1972) the tibe
Signodontini, of fine murid subfamily Cricetinae, distinguished by the
Possession of a complicated glans penis with a finree digitated baculum,

 

30

is essentially Soufin American and probably originated in boreal America,
from which it spread into Eurasia and South America. On fine other hand,
fine Peromyscini, characterized by a simple glans penis, with an unbranched
and normally elongated baculun, is essentially North American and
apparently replaced fine Sigmdont'ni , when finey probably disappeared
from boreal America during Oligocene. The conplex—penis type Cricetinae
seem to have more recently invaded Cental and Norfin America. This
hypothesis appears to be supported by sore ectoparasites, such as fleas,
occurring on these mammals. The flea genre Polygeiis, whidn has moved
into Middle America wifin its complex-penis type Cricetinae and
Caviolorphhosts fronSouthAmerica, is agoodindioatorof suchan
event (Wenzel and Tipton, 1967).

InSouthAmerioa, aswellasinofinerareas oftheworld, fineearly
history of bats is obscure because of fine paucity of fossils. However,
it is estimated finat finey probably appeared in finis continent after fine
Paleocene (Patterson and Pascual (1.29. 99$”) Hershkovitz (1969) believes
finat bats vere probably well established in north-western South America
and fine Isfinmian region during most of fine Tertiary. Storms may have
played an important role in the interchange of bats between the Americas
when fiey were separated.

During Pleistocene times, South America received from North America
a diversity of manuals such as horses, mastodonts, tapirs, peccaries,
camels, deer, some cricetine rodents, squirrels, rabitts, canids, bears
and other carnivores . The fauna interchange between North and South
AmericaduringfinelateTerliary, aswellasfineradiationofmany
mamalgronpshasbeenfinesrbjectofmehspeculatonandeventodayro

31

carnal explanation is available. Ibwever, it appears finat fine majority
of fine families and genera of South American mammals originated on finis
continent when it was conpletely or partially isolated.

At fine present time fine northern sector of South America contains
fine major portion of the marmalian fauna, while fine southern sector is
exceedingly poor in diversity and mnmber of species (Fittkau, 1969;
Osgood, 19%). Information based on Baker (1967), Cabrera and Yepes
(19m) and ofiner aufinors, as well as on personal data, reveals that the
major portion of soufimestern Colombia, corresponding to Pacific Coastal
Lowlands , maintains a nurber of ubiquitions mammal species in addition
to finose forms possessing a more restricted distribution. The
remaining mammalian fauna of fine southwest region of Colombia is
fundamntally represented by a limited number of Andean elements .

The recent biogeographical. approach for fine Neotopical region
presented by Muller (1973) is followed for the brief zoogeographic
chscription of mammals of the Departamento del Valle and the rest of
southwestern Colt-bis. miller based his interpretation of
zoogeographical patterns on dispersal centers devised from comparative
studies of plant and animal distribution .

Colombia and several ofinern territories of topical America belong
to fine Brazilian Subregion of fine Neotopical Region (Hershkovitz, 1958).
The complexity of ecological situations in finis ration has provided for
fine existence of diversified mammalian fauna. Many areas, particularly
finose humid zones rich in vegetation and not yet disturbed by man,
offer optimal condition for sore populations of manuals , especially
rodents and bats. They possess good reproductive potential and have been

 

32

able to occupy a number of ecological niches through the development

of specialized features. Those lands characterized by drier conditions
and a low proclucdvity of plants, particularly of fnnit trees, as well.
as finose sonnewhat disturbed by man, shot, as a rule, less diversity and
smaller populations of male. In a broad sense, sylvan species are rot
too selective in fineir' food and habitat requirements , while pastoral

7 species seen to display a higher level of adaptation by their food habits
and locomotor organs.

Among the Neotopical dispersal centers outlined by MUller, the
following directly concern the Departamento del Valle and ofiner major
portions of southwestern Colombia: 1. The Cauca Center, 2. The
Colombian Montae Forest Center, 3. The Colorbian Pacific Center, and
In. The North Andean Center. For this discussion it is convenient to
follow finese zoogeogaphic divisions . Figure 5 has been prepared using
as a source the map given by Miller (1973).

The Cauca Center. As defined by Miller, this zone is represented
by the Cauca and Patia valleys, which lie between the west and cental
Cordilleras of Northern Colonbia, and are separated from each other by
fine Popayan plateau at an elevation of 1750 meters. Like many other
areas of Colombia, finis territory is being progressively deforested;
however, it still. harbors many mamalian species, sore of which are
widely distributed in other areas of fine Continent. Some of fine faunal
elements finat previously existed in finis territory have probably
disappeared many years ago. Paunal indicators of finis zone are fine
woolly opossum, Calnn'onys derbianus derbianus , and fine red squirrel;

Sciurus granatensis valdiviae, which are found in the Cauca and Patia

Io.

 

0“.

‘kn.

..n. at .....u. .o i 3. .. a in... m..." man. “new are are . he saw he at

n 1 ‘h|

. . , . H. . .
.
e .
e n , n n
o . .
.
a
.- V
n
o
, . . o
n .
. .n
i n .
.
O I
. n u
. v
. r
o
i
Q I
of. n a
O O
o.
O
. s e
n n
_
n O a
I v
y
n n
a . e a
. .
o a s
c D u
o O
W 0
a n n o
O
. .n
w v n n I n .n .
O .
O
I i C
a e
_
0
n v
.
h I O 0
~ . I
.
I
O n v
1
\ o Q a
a n v I p O
I i
r O
. r .
U
I .'
4 4
. I
. V O \
u ‘n a a
U '
u n .
I O
n . 0
e I
. . i
I.
e o ,
., .0.

.r,.3. /

‘ n
\l;‘

33

valleys. 'Ihe ancestors of these manuals evidently care from the north.
According to Miiller, fine slight differentiation between fine populations
of finese subspecies in the two valleys suggests that fine Popayan plateau
is not a stong barrier to species adapted to open habitats . In addition,
fine forest bias of the southern part of the Cauca Valley did not prevent
a spread of fauna during post-glacial times .

Available information indicates that South America was under fine
inflnence of drastic climatic changes during the Quaternary (Hammen,
1961; Patterson and Pascual, 1963; Vanzolini, 1973). Haffer (1967)
considers finat climatic changes during fine Pleistocene and post-Pleistocene
had more influence finan orogenic events on fine fauna west of the Andes.
He also states finat isolation and differentiation of most Pacific species
occurred after fine early Pleistocene uplift of fine Northern Andes ,
probably under orographic conditions similar to finose existing today.

Some ofine fauna elements of the Cauca Center are fine marsupials
Didelgn_1§' marsupialis marsugialis, Chironectes minimus, Marmosa impavida
aging, Hnilander opossum gisescens, fine rodents m alfaroi
M, Hem austalis austalis , _S_i_godon hispidus 1_>_ogotensis,
W Manda brumnens, and Agouti. paca guanta, fine bats
Artibeus jamaicencis , Pteropteyx kaplei lgplei , Molossus molossus
@193, fine edentate Dasypus novemcinctus and carnivores such as
@cyon cinereoargenteus , Mustela frenata, Potos flavus megalotus and
Elam} and fine brocket deerMazama americana zetti.
PEColonbian Montane Forest Center. This dispesal center seens to be
confined to fine foresthiores of Colombia (with the exception of the
Sierra Nevada de Santa Marta), Ecuador and Venezuela. It is divided

3n

into to subcenters, namely, the West Andean and the East Andean.
Inasnmnch as affinities have been found between fine animal populations

of fine Cental Andes and finose of fine West Andean Subcenter, fine Cental
Andes is included with fine West Andean Subcenter (Miiller, 10c. cit. ) .

. 'Ihe fauna of the Colonbian Montane Forest Center may be cornsideed
subtropical and displays similarities wifin those of fine Colorbian Pacific
Center, which is essentially topical. This condition is explained by
fine fact finat most of fine species of fine Colombian Montane Forest Center
have originally evolved from ancestors that came from the lower lancb .

Amang fine most characteristic manuals of the Colombian Montane
Forest Center are: Didelphis azarae, Metachirus nudicaudatus
colomnbianus, Philander Opossum grisescens, Eptesicus brasiliensis andinus,
M calig'nosus monticola, Ms munchiquensis, Nectonys alfari
esmeraldarum, Rhig‘donyg latimanus similis , Thomasomy_s_ aureus popayanus ,
Reifinrodontornys mexicanus milleri , I_ch_fi_nyg_ry_s midrobates nicefori ,
Sciur'us Meanii caucesis, Echinoprocta rufescens, Dasyprocta
fuliginosa oandelensis, m brasiliensis fulvescens, Nasua nasua
oandaoe, mg finonasi medellinius, and Aotus tivirgatus lenurinus.
'Ihe Colonbian Pacific 993g. The extensive territory of lowlands in the
western part of Colonbia involves the major portion of the center. This
territory continues to fine north along fine base of the mountains and ends
on lands wateed by the Magdalena River. To the soufin, finis center
encroadnes upon a portion of fine rnorthern part of Ecuador. This center
is fnrrthe divided into two subcentes: fine Nechi, which encompasses
theareabetweenfineRio Sinuandfine lowerreaches offine Rio Cauca,
and the Choco, which includes fine area west of fine Andes.

 

I i

35

It is possible that during the Tertiary, fine gap separating Cental
and South America divided the territory row lcnown as the Colombian Pacific
Center. This finesis stems from the existence at fiat time, of a seaway
south of Panama and fine Gulf of Uraba, which connected fine Caribbean Sea
and fine Pacific Choco basin of Western Colombia (Haffer, 1967). This
seawaywas closeddur-ingthe late Pliocene andseems tocorrespondto
fine Bolivar Geosyncline discnssed by Nygren (1950) and Hershkovitz (1968).

The glacial and interglacial periods of fine Pleistocene evidently
influenced fine Pacific lowlands of Colombia. Glaciation of the
mountains produced considerable temperature reduction and high humidity.
At finis time, fine sea level lowered about 100 m and extensive movements
of fauna occurred between fine Amazonian region and fine tens-Andean
area of western Oolonbia and Cental America. The integlacial peiods
wee particularly drien in the northern part of Colombia, when the humid
forest moved soufiward due to fine influence of winds, and fine sea level
roseabout 30toSO m. Asaresultoffinis conditionfineMaraoaibobasin
and ofiner parts of fine Northern Colombian plains wee flooded (Haffer,
£03. 992).

Studies made on birch, lizards and amphibians presently inhabiting
this cente, have indicated fine origin and dist-ibution patterns of sore
of finese vertebrates. A stong relationship of the Pacific lowland fauna
of Colonbia to finat of fine Amazon region suggests fine interchange of
animals during finose remote times of fine Pleistocene. Today, the Pacific
mamlian fauna of the Departamentos del Valle, Onoco, Cauca and Narifio,
inhabiting fine dense forests of the coastal lands, are, wifin sore
exceptions, similar to finose of fine Pacific lands of Easten Panama

36

and Northern Ecuador.

Sane elements of the mammal fauna of the Colombian Pacific Center
are Philander ogossum melanurus, Marmosa robinsoni, Oryzom capito,
m tenuipes tenuipes, Proechimys senispinosus, Proechinys
guyanensis, Hoplorys gymnurus, Heterogys austrelis, Microsciurus
flaviventer isthmius, Dasyprocta punctata chocoensis, Hydrochaeris,

Uroderma bilcbatun bilobatum, Chiroderma villcsum, Vampgessa Lnymphaea,

 

Carollia perspicillata, Clossophaga soricina, Artibeus cinereus ,

 

W helleri, y_. dorsalis, N0ctilio labialis labialis, Desmodus

rotundus, Molcssus molossus, Cebus cgucinus, Bredipus griseus,

 

 

Chologpus hoffmanni, Potos flavus, Bassaricyon gabbi, Procyon

 

 

cancrim panamensis, Felis onca, Felis pardalis, Galyctis allamandi,
‘I‘ggsu gcari, TaLassu taiacu, Mazama americana, Tapirus bairdii.
_‘ILne; m Andean Center. This area is the Central and Eastern Cordillera I
in Colombia and the Andean mountains in Ecuador and Peru. The most
dnaracteristic bicme of finis center is fine paramos with its selected
plant and animal life. To this center belong such mammals as the
spectacled bear, 'I‘renamtos ornatus , Pug mephistogniles , Sylvilagus
brasiliensis andinus, Nasuella clivacea, Caenolestes fuliginosus,
Marmsa $32.54 and 'Ihcmnasonys cinereiventer cinereiventer.

According to Minller, it is apparent finat 28 species of the
37 North Andean faunal elements (mere fimn 75%) belong to families
of North and Central American origin.

Chapman (1917), refering to birch from fine panama zone, indicated
that fine maj crity were derived from the sea level eqm‘valent of this
zone in soufinern South America. It sears logical to assure finat the

37

same circmnstance occurred with the manuals also.

 

38

 

    

.3...

a;

a?:

9’
.L%

 

 

:5
. _ _ ___::_:__..
___:________:_________

 

cw nouns: ”I “m

 

THE MIA” PACIFIC CENTE.

THE

T"; c on can:

  

 

Fig, 5 .

 

 

PACIFIC OCEAN
ICuADOn

 

 

 

ACCOUNTS OF FLEAS OF SOUIHWESTERN COLOMBIA

In finis presentation taxonomic descriptions are limited to those
forms that are either new to science or have been previously described
from one sex. Only the citation of the original description for every
taxon is given. For complete syrnonymny , type data and other distribu-
tional records fine reader is referred to Johnson (1957), Tipton and
Méndez (1966) and Tipton and Machado—Allison (1972). The principal
sources of information of Colorbian manual hosts have been Allen (1912 ,
1913, 1915, 1916), Borr'em (1967), Cabrera (1958 and 1961), Cabrera and
Yepes (1990), Her'smcovitz (191+1, 19'47, 19'48, 19ln9, 1960, 1962),
Gyldenstolpe (1932), Tate (1932a,b, 1935), and Osgood (1912).

The following report pertains to taxa from southwestern Colombia
available to us for study and forms finat are not represented in our
material but have been reported before from that territory or are likely
to occur finere. The order followed here is that of Johnson (1957).

SUPERFAMILY CERATOPHYIIDIDEA
FAMILY HYS'I'RICHOPSYILIDAE
SUBFAMILY CI'ENOPIW
TRIBE WRATINI
Neotyphloceras rosenbergi (Rothschild)

(Figure 8)

39

1+0

Typhloceras rosenbergi Rothschild, 190'4, Novit. 2001., 11:639, P1.
13, Figs. 68-69; Pl. lu, figs. 71, 7a.

Material examined: Ex Didelphis marsupialis, (71-621), 1 d',
Pichindé, 1600 m, Depto. del Valle, 6.VIII.1971, H. Tyndale-Biscoe;
(71-627), 1 a", same locality, date and collector; ex Didelphis
m, (00110), 1 d', Finca Holanda (near Péram de Chinche), 2700 m,
Depto. del Valle, 28.X.1975, E. Mendez 6 L. Velasquez; ex m
caJiginosus (Hm-nus), 1 9, Quebrada Honda near Pichindé, 1800 m,
Hmicipio de Cali, Depto. del Valle, 13.VIII.1965, H. Trapido; (Hm-208),
l 9, same locality, 8.3.1965, H. Trapido; (Hm-385), 1 a", same locality,
lll.XI.1965, H. Trapido; (HIE—1260), 1 9, same locality, date and
collector; (Hm-lugs), l 0", same locality, 15.111.1957, H. Trapido;
(HBO-1623), l o", sane locality and collector, 29.111.1957; (HIE-1631+),
l 9, same locality and collector, 31.111.1957; (Hm-2192), 1 0'2 9,
Pichindé, 1900 m, Murnicipio de Cali, Depto. del Valle, 27.1.1968;
(00007), 2 o’l 9, Pichindé, 1900, Municipio de Cali, Depto. del Valle,
21-25.I.19714, M. Thom 6 L. Velasquez; ex Omcmys alfaroi, (HIE-209),

 

l 9, Quebrada Honda near Pichindé, 1800 mn, Mnmnicipio de Cali, 5.1x.1955,
H. Trapido; (HTc-lugl), 1 9, same locality and collector, 15.111.1957;
(MI‘ uzss), l 9, Florida, 8 km s. B. " La Diana," 1700 m, 3.x.197u;

ex cryzcmys albigularis (Hm-1237), l c', Valle del Rio Pichindé,
1700—1900 In, Municipio de Cali, Depto. del Valle, 3l.X.1966, H.

‘I‘rapido; (RFC-1308), 2 do: sane locality and collector, 13.XII.1966;
(Hm-1381), l 9, sane locality and collector, 13.XII.1967;

(ETC-1382), l c', same locality and collector, 11.x.1957; (Hm-1385),

'41

l 9, same locality and collector, 12.1.1957; (Hm-1835), l o", Cerro
Munchique, 60 km by road west of Popayan, Pefia del Perm, 2160 mn,
ll.V.1967, H. Trapido; (RFC-18146), l a", same date, locality and
collector; (mt-18w), l 9, Cerro Munchique, Finca El Retiro, 2200 m,
Depto. del Cauca, 12.V.1967, H. Trapido; (Hm-2771), l 9, Finca la
Flora, Quebrada Norte, Pichindé, 1900 m, Depto. del Valle, 1.VIII.1968,
H. Trapicb; (00065), 5 d', Saladito, km 12, 2000 mn, Depto. del Valle,
17.III.197H, E. Mendez 8 L. Velasquez; (00105), 3 c", Finca Holanda
(near Pareto de Chinche) 2700 m, Depto. del Valle, 28.X.l975, same
collectors, (00108), 1 9, sane locality, date and collectors; (00109),
2 9, sane locality, date and collectors, (00112), 1 9, same locality
and collectors, 29.x.1975; (00121), 1 6'1 9, same locality and
collectors, 30.x.1975; ex Oryzomys (Oligcryzouys) sp. 1 9, Quebrada
anda near Pichindé, 1800 m, Hmicipio de Cali, Depto. del Valle,
2.x.1955; ex Rhipidomnys latimanus, (Hm-337), l 9, sane locality,
25.X.1965, H. Trapido; (ETC-871), 1 d’, Valle del Rio Pichindé,
1700-1900 111, Municipio de Cali, Depto. del Valle, 12.VII.1966, H.
Trapicb; (nut-1295), 1 9, sane locality and collector, 8.XII.1966;
(HI‘C-llnl'l), 1 9, same locality and collector, 12.VII.1966, H.
Trapicb; ex Rhipidonfi similis, (HID-1805), l 9, Cerro Munchique

(60 Joe by road west of Popayan, sitio No. 1, 2500 m, Depto. del
Cauca, u.V.1967, H. Tmpido; ex Thonasomys m, (71-561), 1 9,
Pilimbala, 3100 111, Depto. del Cauca, 29.V.1971, H. Tyndale-Biscoe;
ex Thanasgys cinereiventer, (Hm-1812), l 9, Cerro Munchique,

‘42

60 km by road west of Popayan, sitio No. l, 2500 m, Depto. del Cauca,
5.V.1967, H. Trapido; (RIC-1828), 1 0", same locality but sitio No. 3,
2500 m, 9.V.1967, H. Trapido, (Hm-2m), l 9, Laguna de La Cocha,
2700 m, Depto. de Narifno, 15.V.1969, H. 'I‘rapido; (Hm-18w),

o", Cerro Munchique, 60 kms by road west of Popayan, Pe‘ia del

Farm, 2160 m, Depto. del Cauca, 12.V.1967, H. Irapido; (ETC—21108),
l d’, Iagma de La Cocha, 2700 m, Depto. de Narif'no, l6.V.l968,

H. Trapido; (RIC-21:15), l 6", same locality and collector, l7.V.1968;
(INC-2MB), l 0’, sale locality, date and collector; (RIC-21122),

2 a", sane locality, date and collector; (BIC-21127), 1 a", same
locality and collector, 18.V.1968; (I'M-2180), 1 0", Km 38 between
Paste and Sibuncby, 3100 m, Conisaria de Putumayo, 22.V.l968, H.
Trapicb; (RIC—211111), 1 0", Km 33 between Pasto and Sibundoy, 2900 mn,
Depto. de Narifno, 22.V.1968, H. Trapido; ex 'Ihonascgys fuscatus,

 

(RIC-11103), 1 o”, Valle del Rio Pichindé, 1700-1900 m, Municipio de
Cali, Depto. del Cauca, 7.XI.1966, H. Trapido; (HIE-11415), l 6‘,

sale locality and collector, 17.1.1967; (RIC-1633), 1 0", same locality
and collector, 31.111.1957; (Hm-1670), 1 9, Pichindé, 1900 m,
Municipio de Cali, Depto. del Valle, 19.V.1967, H. Tralido; (HIE-1673),
l o‘, sane locality and collector, 23.V.1967; (HIE-1676), 1 d", sane
locality, date and collector; (REC-1688), l 9, same locality and
collector, 26.V.1967; (HID-21102), l 9, Laguna de La Cocha, 2700 m,
Depto. de Narifio, lS.V.1968, H. 'I‘rapido; (RFC—2955), 1 6', Rinc5n

del Yarural, Pichindé, Depto. del Valle, 8.V.1969, H. Trapido;
(Hm-2978), 1 6'1 9, same locality and collector, 28.V.1968;

H3

(HIE-32145), l d', Finca La Flora, Pichindé, Depto. del Valle,
26.VII.1969, H. Trapido; (00050), 2 9, Saladito, Km 12, 2000 m,
Depto. del Valle, lS.II.197l+, E. Méndez 8 L. Velésquez; (00051),
2 Ju 9, sane locality, date and collector; (0005M), 2 6', same
locality and collectors, 16.11.197I4; (00071), 1 c”, sane locality
and collectors, 18.11.197u; (00075), 1 o", 2 9, sane locality and
collectors, 19.II.197Q.

Remarks. In addition to Colonnbia, this species is
distributed in Venezuela, Ecuador and Perfi. It is regularly fomnd
on a variety of rodents and less conmnly on marsupials. Our
material contains specimens from Didelphis azarae, Q.
marsgialis, Oryzcmys (Oligoryzorys sp.), _O__._ (Oryzonmys albigularis,
_Q. (Orywnys) alfaroi, _O_. (Melanomys) calig'nosus, Rhypidorys
latimanus, 3. similis, Thonasonys aureus, _T_. cinereiventer,

_T_. fuscatus and Thonasonys sp. Other hosts recorded in the
literature for this flea are fine following: Phylander oEsum,
Marmesa, m, cram, Rheomys, Stictomys, and gm. The
variety of hosts used for this flea certainly indicates a low

degree of host specificity.

an

TRIBE AIDRATOPSYILINI

Adoratppsglh (Tritgpsylla) intermedia copha

 

(Figure 9)

Stenopsylla intermedia copha Jordan, 1925, Novit. 2001., 33:391,
fig. 15.

Material examined: Ex Didelphis azarae, 1 5'1 9, vicinity of Cali,
Depto. del Valle, 11.x.1959, v. E. Thatcher; 2 5‘2 9, same locality and
collector, 16.X.1968; Didelphis marsupialis, (ETC-2661+), 1 6'1 9, La
Maria, 1900, Municipio de La Cumnbre, Depto. del Valle, 3.XI.1967, H.
Trapicb; (71-521), 1 61 9, Pichlindé, 1500 mn, Depto. del Valle,
5.VIII.1971, H. Tyndale-Biscoe; (71-525), 2 5’2 9, sane locality, date
and collector; (71-525); 1 6'1 9, sane locality, date and collector;
(71-527), 3 6'3 9, same locality, date and collector; (00091), 1 o”

5 9, Iago Calima, 1950 m, Depto. del Valle, 23.11.197u, B. Mendez 5

L. Velasquez; (00092), 2 o' 3 9, sane locality, date and collector;
(00093), 7 6'7 9, same locality, date and collector; ex Philander
mg, (71-523), Pichindé, 1500 111, Depto. del Valle, 6.VIII.1971,

H. Tyndale-Biscoe; (00039), 1 9, Alto Anchicaye’n, 550 m, Depto. del
Valle, 11.11.197u, E. Méndez s L. Velésquez; (000u3), 2 9, sane
locality, date and collectors; ex Oryzomys caliginosus, l d; Pichindé,
1600 mn, Depto. del Valle, 21-25.I.197ln, M. Thomas 8 L. Velésquez;

N»,

#5

ex Thonascmys fuscatus, 2 0’3 9, Saladito (lfin 12), 2000 m, Depto. del

 

Valle, 15.11.1979, B. Mendez 5 L. Velasquez.

Remarks. This subspecies has been reported from Colombia,
Panama, Ecuador and PerCn, from sea level to over 3000 meters. It is
a common parasite of marsupials and sonetimes display a high degree
of infestation on a single host animal (Tipton and Mendez, 1966).

In southwestern Colonbia it has been obtained from Didelphis azarae,
Q. marsupialis, Metachirus nudicaudatus, Philander opossum and
'I‘honasgnfi fuscatus. Elsewhere this flea has been also recorded from
Oryzom caliginosus and Proechilnys semispinosus.

IHWUIR'PYGIOPSYLLIDAE
SUBEAMIEY PYGIOPSILLINAE
Ctenidiosomus re_x_ Johnson

Ctenidicsonnus rex Johnson, 1957, Men. Ent. Soc. hash. 5:50,
P1. 20.

Remarks. The type material (2 males and 2 females) from San
Agustin, Departamento de Huila, Colonbia, represents the only specimens
know of this taxon. These specimens were collected from Thomasonys,

m and M. It seems probable finat a species of
‘Ihomaggnys (probably I, laniger , is fine natural host of finis flea.

Ctenidioscmnus traubi Johnson
(Figure 10)

Ctenidiosonmls taubi Johnson, 1957, Men. Ent. Soc. Wash. 5:ln9-50,

P1. 17; P1. 18, fig. 5; P1. 19, figs. 1, 2).

146

The original description is based on the holotype female , ex
Caenolestes obscurus, Colombia, Depto. de Antioquia, Sansc’rn, 7 km. E.
of Péruo, 3,160 m., 18 Oct. 1950, P. Hershkovitz collector. The male
is described helm.

Material examined: ex Caenolestes obscurus, (71-521), 1 9,
Puracé Park, 3500 m, Depto. del Cauca, 30.V.1971, H. Tyndale-Biscoe;
ex Mm; (71-561), 1 d; Pililnbala, 3100 m. Depto. del Cauca,
29.V.1971, H. 'I’yndale-Biscoe; ex Thonasomys cirnereiventer (HI‘C-2l156),
l o", Comisaria Putunayo, K 77 between Sibundoy a Mocoa, 2200 m,
27.V.1968, H. Trapido.

DESCRIPTION OF MALE.

_iI-had; (Fig. 10 A): Strongly fracticipit, with frons evenly rounded,
devoid of clypeal tubercle . Preantennal region provided wifin 3
distinct discs and abundant nicropores; principal bristles of preantennal
region arranged in 2 rows , secondary bristles very shot, scattered
mainly on preocular area. Bye deeply excised ventrally, weakly
pignnented. Postocular area wifin 2 pits located just below eye. Genal
area bilcbed, its anterior lobe or genal process broadly rounded, not
accnminate. Posterior lobe of gena evenly rounded, moderately broad.
Postantennal region characterized by anterior micropores, several pits
profneely distributed and 2 rows of bristles in addition to short
bristles distributed on dorsal marg‘n and in front of antenna, mainly
on anteual fossa. Pedicellus of antenna covered wifin short bristles.

m: Very setose. Pronotum having 2 rows of bristles preceding
conbofabout 26 spines. Mesonotunat leastwith 3well definedrows of

147

bristles , remaining bristles short , concentrated on anterior pronotal
region. Mesepisternum wifin few non prominent bristles located on
anterocbrsal area near pleural ridge of mesofinorax. Mesepimere with
bristles of different sizes. Metanotunn covered with about 3 or In
defined rows of bristles , posteriormost row combines long bristles and
short interoalaries. Ofiner bristles not arranged in rows belong in
anterior metanotal area. lateral metanotal area apparently with no more
than 1 bristle . Metepisternum with oval outline interrupted by anterior
projection of metasternum, provided with few bristles . Metepimere with
3 rows of uneven bristles.

Abdonen: Connbs present on terga II—V, armed wifin variable nunber
of spines (in the two specimens examined), respectively, 114-15; 114-115;
12-13; 111-15. Upper antesensilial bristle about 1/2 as long as lower
bristle.

Modified abdonninal EM: Eight ter'gun reduced to
subtriangular plate provided with broad spiracle, having goup of short
bristles near antesensilial bristles. Bight sternum large, ensheating
principal structures of genitalia, provided with nunerous marginal and
inner bristles, its caudal margin entire, wifinout sinus. Clasper large,
sonewhat pyriform, projected anteriorly into short mnanubrium curved
(inwards. Process provided wifin subrounded apical expansion,
largely squamose and bearing group of bristles distributed on outer and
inner surfaces. Posterocaudal margin of process armed with four long
bristles. vaable process of Clasper broadest near its basal area,
becoming narrower apically, showing bristles of variable size. Ninfin
sternum (Fig. 10 B) resembling those of other species of gene. Distal

us

arm club—shaped, having broad and rounded apex armed with In stout
bristles on caudal margin; remaining bristles smaller, scattered over
most of arm. Prondmal arm of ninth sternum narrow at base, broad at
apex. Aedeagus resembles that of Ctenidiosomus perplexus Tipton and
Machado-Allison. Median dorsal lobe broad, with rounded dorsal margin,
producing caudal subacunminate blade. Lateral lobes narrower finan
distal lobe, sligntly arched. Crochets crescent shaped. Aedeagal
apodemal rod arched apically. Penis rods strongly coiled, extensively
fimbriate on apical portion.

Remarks. Up to fine present tn'lne, Ctenidiosomus traubi is lonown
only from Cololbia, where it has been collected in localities with
elevation ranging from 2200 m to 3500 m. The scant material available
has been collected on Caenolestes obscur-us, M m and '_I'_.
cinereiventer. As Lewis (197%) has pointed out, it is likely fiat fine
preferred hosts of Q. _t_r__'a_ub_i are rodents.

FAMILY SI'EPHANOCIRCIDAE
SUBFAMILY CRANIDPSYIIINAE
TRIBE CRANIDPSYLLINI
Cleopsylla monticola Suit
(Figure 11)

Cleopsylla mnontioola Suit, 1953, Bull. Brit. Mus. (Nat. Hist.)
Eh‘tOlDL, 3 (5):193, figs. 13, 15, 17, 19, 20.

Material examined: Bx 931mm, (Hm-1505) l 9, Cerro
Munchique, 60 km by road west of Popayan, Sitio No. 1, 2500 m, Depto.
(bl Gama, '4.V.1967, H. Trapicb; (RIC-1817), 1 d", sane locality and

‘49

collector, 5.V.1967; (001014), 2 d‘, Pinca Holanda (near Parano de Chinche),
2700 m, Depto. del Valle, 28.X.l975, E. Méndez and L. Velasquez; (00105),
1 6'1 9, sane locality, date and collector; (00106), 2 0’: same locality,
date and collector; (00107), 2 o", sane locality, date and collector;
(00112), 1 6; sane locality, date and collector, (001l5), 1 a", sane
locality and collector 29.X.l975; (00117), 1 a", sane locality and
collectors, 30.x.l975; (00118), 1 9, same locality, date and collectors;
(00119), 2 d'2 9, sane locality, date and collectors; (00121), 1 6'2 9,
same locality, date and collector; ex Rhipidonys similis, (HIE-1805),
1 d', sane locality and collector, n.v.l957; (Hm-1833), 1 5’1 9, sane
locality and collector, Sitio No. 3, 2500 m, 10.V.1967; ex Thomasonys
cinereiventer, (HIE-21113), l 6', Laguna de La Cocha, 2700 m, Depto. de
Narifio , 16.V.1968, H. Trapicb; (RFC-21418), l 9, same locality and
collector, l7.V.l968; (RFC-21407), l 0", same locality and collector,
16.V.1968, ex ‘Ihomasw fuscatus, (ETC-1690), 1 d; Pichindé, Finca La
Horn, 1900 m, Depto. del Valle, 30.V.1967, I'LTrapid).

Remarks. Reports of finis helmet flea are from Ecuador, Colonbia
and Veneznela. In Colombia, it has not been collected above 2700
meters elevation. The range of vertical distribution for finis species
in Venezuela is from 120 meters to 11m meters (Tipton and Machado—
Allison, 1971). The following mammals have been found harboring Q.
monticola in soufinrestern Colonbia: gyms albigulg’s , Rhipidonys
similis, W cinereiventer, L fuscatus, and W sp.
Other hosts recorded in fine literature are Caenolestes fuliginosus,
Ilidelphis marsupialis, Marmosa fuscata, M. dryas, Oryzcms minutus,
Mvemstus, Ma” 'Ihonasomys hylophilus, 11;. lani er,’

50

T. vestibus, Chilgyg instans and birds.

§phinct0psylla diomedes Johnson
(Figure 12)

Sphinctqpsyfla dionedes Johnson, 1957, Man. Ent. Soc. Wash.
5:68, P1. 32. '

This species was originally described from two male specimens
ex Caenolestes obscurus, Colombia: Depto. of Huila, San Agustin, San
Antonio, left bank of Rio Magdalena (Cordillera Central), 2200 m,

214 Aug. 1950. F. Hershkovitz collector.

Material examined: ex Caenolestes obscurus, (71-519), 1 o",
Putnacé Park, 3500 m, Depto. del Cauca, 29.IV.197l, HTyndale-Biscoe;
(71-521), 2 5’3 9, sane locality and collector, 30.IV.1971; (71-539),
1 8'1 9, sane locality and collector, l2.V.l97l; (71.5111), 1 0‘1 9,
sane locality and collector, lu.v.l971; (71-579), 3 6'2 9, sane
locality and collector, 2h.VI.l97l, (71-580), same locality and
collector, 2u.VI.1971; 1 9 (REM No. 3755), El Soche, 2700 m, Municipio
de Soacha, Depto. de Cundinamnaroa, Colonbia.

26.IX.1969, R. B. Mackenzie.

WION OF M.

Head (Fig. 12 A): Frans mgin moderately rounded. Helmet comb
consistingof13 spines. Genalcomboszpines.
'Ihcrex: Pmnotum (Fig. 12 A) having 2 rows of bristles and

 

conspictous comb of 9 spines per side. Remaining thoracic structures

51

and legs as in male.

Abdomen: Terga I-IV bearing apical spineletes . All terga wifin
two rows of bristles but anteriormost row more reduced. Tergun VII
provided wifin 2 subequal antesensilial bristles.

Ibdified abdominal segments (Fig. 12 B): Posterior margin of
tergun VIII sinuous, having 2 groups of large, stout spiniformn bristles,
upper group consists of 9-6 bristles, lower group of 1-3. Both groups
preceded by scattered bristles of variable size and location, in
addition to short and moderate size marginal bristles . Spiracle of
ter-ganIIIwithverybroadbasal portion. Sterna II-VIwifin lrow
of 6 bristles. Sternum VII bearing group of 3-11 stout spiniformn
bristles on each side, in addition to several inconspicnous bristles.
Posterior margin of finis sternum almost straight , not indented.
Sensiliun with about 11 sensory pits per side. Dorsal anal lobe
armed with several bristles. Ventral anal lobe wifin only 1 or 2
bristles . Anal stylet short and stout, cbrsally and ventrally convex,
its apical bristle about twice fine lengfin of stylet body, accompanied
by minute ventral bristle and nasal bristle of moderate size.
Spermatheca (Figs. 12 B, 12 C) with divided bulga, having anterior
section globular, fairly reticulate, and posterior section not reticulate ,
followed by short, upturned unpignerrted hilla. Main body of bursa
copulatrix short, sinnous, weade sclerotized.

Remarks. This species is endemic to southwestern Colonbia where
it has been found in areas between 2200 and 3500 meters. All specimens
presently existing in collections are from Caenolestes obscurus which
seems to be the natural host.

52

Sphinctcpsylla tolmera (Jordan)

Craneopsylla tolnera Jordan, 1931, Novit. 2001., 36:3lu, fig. 5.
Material examined: Bx Thonasonys cinereiventer, (HI'C—Zuou), l 9,

 

 

Laguna de La Cocha, 2700 m, Depto. de Narifno, 16.V.1967, H. 'I‘rapido;
(Inc-21.33), l 6‘, same locality and collector; (colon), 1 d‘, Finca Holanda
(Near Paramo de Chinche), 2700 m, Depto. del Valle, 28.X.1975, E. Méndez
8 L. Velasquez.

Renar'ks. The geographical range of g. tolmera involves Colonbia,
Ecuador and Venezuela in some areas exceeding 2000 meters elevation.
Rodents of the gems 'Ihorasonys probably are the preferred hosts in
Colonbia and Ecuador. In Venezuela this flea seems to be more associated
with Oryzonys minutus. Of 36 Hales and 76 females recorded by Tipton and
Madnacb—Allison (1972), 32 males and 63 fenales were recovered from #6
specinms of this rodent.

Plocopsylla plyllisae Suit
(Figure 13)

Plocopsylla phyllisae Smit, 1953, Bull. Brit, Mus. (Nat. Hist.)
Eh’aonoL, 3:197, figs. 25, 25, 23, 30.

Material entwined: a: Caenolestes obscums. (71-521), 2 6' 3 9,
Puracé Park, 3500 m, Depto. del Cauca, 30.IV.1971, H. 'I‘yndale-Biscoe;
(man), 3 9, same locality and collector, lu.v.197l; (71-557), 2 cf's 9,
sane locality and collector, 28.V.197l; (71-579), 3 6', same locality and
collector, 2u.v1.1971; (Rm 3765), 2 9, Bl Soche, Mnrnicipio de Soacha,

53

2700 m, Depto. de Cundinamarca, 26.3.1969, R. B. Mackenzie.

Renarks. This species is confined to some territories over 3000
meters elevation in Ecnador and Colonbia. The holotype male was secured
from m sp., nevertheless, the material (11 defend 21 99) available
to us for study came from six specimens of Caenolestes obscurus, which
is probably the preferred host. This interesting terrestrial marsupial
lives in dark damp forests of péreros and is crepuscular or nocturnal.

It seem to be prinarily insectivorous (Osgood, 1921; Tate, 1931).

Plocopsylla thcr Johnson
(Figure ll!)

WMJOIWm, 1957, Men. Ent. Soc. Wash., 5:73—7u,
P1. 38 (figs. 1, 2, 3, 6, 7), P1. 39 (figs. 1, 2, 3).

Material examined: Bx Thonasonys cinereiventer, (HIE-1827),

l 9, Ceao Mmchique, 50 km by road west of Papayén, Sitio No. 3,
2500 m, Depto. del Cauca, 9.V.1967, H. Trapido; CHIC-2H0”, 1 d:
Iagma de La Cocha, 2700 m, Depto. de Narifio, 16.V.1968, H. Trapido;
(Inc-2MB), l 9, sane locality and collector, 18.V.l968, (Hrc-2u32),
l a; sane locality and collector, 19.V.1968; (RIC-2H35), 2 9, same
locality, date and collector.

Renarks. Plocopsylla thcr seems to be restricted to some areas
of high elevations (particularly between 2000 and 3000 meters) in
Colonbia. It has been found associated with the crice'dne mdents
m (M) albigularis and Thonasw spp. The typical host
for this flea is probably 'Ihcuasonnys cinereiventer.

5’4

FAMILY ISCHNOPSYLLIDAE
SUBFAMILY ISCHNOPSYLLINAE
Sternopsylla distincta speciosa Johnson

(Figure 15)

Sternopsylla distincta speciosa Johnson, 1957, Men. Ent. Soc.
Wash. 5:100; P1. 148, figs. 3, u; P150, figs. 3, 8.

Remarks. The description of this subspecies is based on a
holotype male, an allotype female, and three female paratypes
ex Tadarida brasiliensis, Peru: Dept. of Cuzco, Quince Mil, 19 June
1950, C. Kalinowski collector. One male and three female paratypes
ex Tadarida sp., Colonbia: Dept. of Huila, Pitalico, 1350 m, 28 Nov.
1951, P. Hershkovitz collector. No other Colombian record are
mentioned in the literature.

FAMILY CERATOPHYLLIM
SUBFAMILY LEPI‘OPSYUINAE
Leptopsylla segnis (Sdnonherr)

(Figure 16)

£13135 £23213 Sdncnher'r, 1811, K. svenska Ventenskakad. Handl.
(2) 32:98, P1. 5, figs. A. B.

Material examined: Ex Rhipidcgys latimanns, (HIE-269), 1 9
merada Honda near Pichindé, 1800 m, Municipio de Cali, Depto. del

55

Valle, 23.3.1965, H. Trapido; ex Rattus rattus, 1 0", same locality

 

and collector, 1.VIII.1965.

Remarks. leptcpsylla segnis is a cosmopolitan species which has
been introduced with the comrensal rodent hosts to many parts of the
world. In South America it seems to be confined to areas of high
elevations in Colombia, Ecuador, PerCn, Venezuela, Brazil, Chile and
Argentina. Typical hosts are various species of Muriche; however,

the true host is the house mouse, Mus musculus.

 

SUBFAMILY CERATOPHYILINAE
Dasypsyllus gllinulae perpinnatus (Baker)

(Figure 17)

Cer’a‘tophyllus perpirmna'tus Baker, 1901+, Proc. U. 8. Nat. Mus.,
27:386, 391, HHS, figs. 1-6.

Material examined: Ex Thomasomys cinereiventer, (ETC—1812),

l o”, Cerro Munchique, 60 km by road west of Popayan, Sitio No. l,
2500 m, Depto. del Cauca, 5.V.l967, H. Trapido; (HIE-1827), 1 o”,
Cerro Munchique, Sitio No. 3, 2500 m, Depto. del Cauca, 9.V.1967,

H. Trapido; (Hm-1853), l 9, Cerro Munchique, Pef'na del Perro, 2160 m,
Depto. del Cauca, 12.V.l967, H. 'I‘rapicb.

Renarks . Dasypsyllus gallinulae perpinnatus is widespread
bird flea which has been recorded from several countries in the New
World. Our specimens represent the first report of this taxon for
Colonbia. It is also known to exist in Canada, the United States of

56

America, Panama, Venezuela and Argentina.

It is interesting to note that our specimens were obtained from
different sites and dates on three Thomasomys cinereiventer. These
rodents were trapped on the ground; however, the fact that they were
parasitized by this bird flea suggests that T_. cinereiventer is
perhaps partly arboreal . The fleas were probably obtained from bird
nests located on trees visited by the rodents. At the present time

little is lcnown about the habits of '_I'_. cinereiventer.

Pleochaetis eggatoris gquatoris (Jordan)

Ceratcphyllus ecmatcris Jordan, 1933, Novit. 2.001., 38:3u|+,
fig. 63, (m).

Material examined: Ex ‘Ihonasonys cinereiventer, (BIC-2926) ,
l 9, Iaguna de La Cocha, 2700 mn, Depto. de Narifio, 18.V.1968,
H. 'I‘rapido; (HI'c-2u55), l 9, Km 77 between Sibundoy and Mocoa, 2200 m,
Comisaria de Putnmnayo, 27.V.1968, H. Trapido.

Remarks . Pleochaetis gguatoris equatoris has been reported from
Peru, Ecuador and Colombia. According to Johnson (1957), it is likely
that specimens from Perfi assigned by Macchiavello (19148) to P.

gallatoris equatoris are P. doles guitanus .

Pleochaetis smiti Johnson

(Figure 18)

57

Pleochaetis smiti Johnson, 1951+, Jour. Wash. Acad. Sci.,

 

ll'+(9):291, 295, figs. 1, 3, 6-8, 10, 12, 13, 16, 21, 25, 26, 31.

Material examined: Ex Caenolestes obscurus, (71-503), 1 9,
Pinacé Park, 3500 m, Depto. del Cauca, 16.IV.1971, H. Tyndale-Biscoe;
ex Oryzomys albigularis, (Hm-1826), l o', Cerro Mundnique, 60 km by
road west of Popayan, Sitio No. 3, 2500 m, 8.V.1967, H. Trapido;
(00108), 1 d’, Finoa I'blanda (near Panama de Chinche), 2700 m, Depto.
del Valle, 28.X.1975, E. Méndez a L. Velasquez; (00115), 1 o‘ l 9,
same host, locality, date and collectors; (00117), 1 9, same host,
locality and collector, 30.x.1975; ex Thomasomys cinereiventer,
(mo-21:32), 1 o”, Laguna de La Cocha, 2700 m, Depto. de Narifn'o,
19.V.1968, H. Trapido; (Hm-2m), 1 9, x. 38, between Pasto and
Sibundoy, 3100 mn, Comisaria de Putumayo, 25.V.1968, H. Trapido;
(mo-2u55), 1 d', K 77, between Sibundoy and Mocoa, 2200 m, Conisaria
de Putumayo, 27.V.1968, H. Trapido.

Remarks. _P. §_m_u._t_i_ is to date lonown from Colonbia, Ecuador
and Venezuela. Judging fron our collection and the data available
in the literature, the vertical distibution of this species extents
from 1980 to 3810 elevation. Tipton and Machado—Allison (1972)
report abundant material (203 males and 208 females) from Venezuela.
Evidence is presented by these authors finat the characteristic host
of P. §l_ni_t_l:_ in Venezuela is OAryzomys mn‘nutus. They recoved 36H
specimens of P. gull fran 158 specimens of this rodent.

I suspect that in Colonbia 2° §_m_ui_ti_ is probably more specific
in gym albigularis; however, the material of this flea that has

,1
On

3.

58

been collected in this country is very scant and does not allow any

final interpretation as to the preferred host .

SUPERFAMILY RHOPALOPSYLLDIDEA
FAMILY RHOPAIDPSYLLIDAE
SUBFAMILY RHOPALOPSYLLINAE
TRIBE PARAPSYLLINI
Tetrapsyllus comis Jordan

(Figure 19)

Tetapsyllus comis Jordan, 1931, Novit. Zool., 37:135, fig. 1.

Material examined: Ex Caenolestes obscurus, (71-522) 2 9,

 

Puracé Park, 3500m, Depto. del Cauca, 1.V.l97l, H. Tyndale-Biscoe.
Remarks . Tetrapsyllus comis was hitherto knom only from

 

Ecuador. Our record represents the first for the Republic of
Colonbia. The scant information existing on this flea does not
allow for the determination of any host preference. We have two
females from Caenolestes obscurus while the Ecuadorian holotype
fenale was taken on Sim sp. It is apparent that 1. gigis
a typical member of the Andean fame of the northwest portion of
South America, perhaps limited in its distribution to Colombia
and Ecuador. The male of this Species remains unknown.

59

TRIBE RHOPALOPSYLLINI

Polygenis bohlsi bohlsi (Wagner)
(Figure 20)

Pulex bohlsi Wagner, 1901, Hor. Soc. Ent. Ross., 35:21,
Pl. 1, fig. 6.

Material examined: Ex mmignflaris, (HID-580), l o",
Valle del 'Rio Pichindé, 1700-1900 m, Depto. del Valle, 19.111.1955,
H. Trapido; (00059), 1 9, Saladito (Km 12), 2000 m, Depto. del Valle,
17.II.197u, E. téndez 8 L. Velasquez; ex Welfaroi, (ETC-677),
5 o"! 9, Sonso, 1000 an, Municipio de Buga, Depto. del Valle, 9.V.1955,
H. Trapicb; ex m caJigincsus, (HTC- 519), 2 6'1 9, Valle del
Rio Pichindé, 1700-1900 mn, Municipio de Cali, Depto. del Valle,
5.Iv.1955; (mo-522), 2 6'2 9, Sonso, 1000 m, Municipio de alga, Depto.
del Valle, III.1955, H. Trapicb; (mo-523), 2 0'1 9, same localtiy,
date and collector; (HID-677), 5 o’" 7 9, same locality and collector,
5.VI.1966; (Hm-750), l 9, Valle del Rio Pichindé, 1700-1900,
Hunicipio de Cali, Depto. del Valle, 23.VI.1966, H. Trapido;
(Hm-579), 2 8’1 9, samne locality and collector, 10.V.1955; CHIC—805),
1 9, same locality and collector, 5.VII.1966; (mt-932), 1 9, Pichindé,
1800-1880 an, Municipio de Cali, VII.1966, H. 'I‘rapido; (Inc-905), 1 9,
Valle del Rio Pichindé, 1700-1900 m, Municipio de Cali, 5.VII.1966,

60

H. Trapicb; (mt-1511), 1 9, same locality and collector, 18.111.1957;
(mo-1259), l 9, same locality and collector, 21.x1.1955; (RIC-2262),

l 9, Pichindé, Mnrncipio de Cali, Depto. del Valle, 9.IV.1968, H. Trapido;
(HID-2957), l 9, Sonso, 1000 m, Mmcipio de Buga, Depto. del Valle,
ll.VI.1958, H. Trapicb; (Hm-2958), 1 6'1 9, same locality, date and
collector; (RIC-2875), l 9, same locality and collector, 12.v1.1958;
(HI'C-2u81), l d; same locality and collector, lu.VI.l958; (Hm-2982),

1 9, sane locality and collector, 15.VI.1958; (HIE—21483), l c", same
locality and collector, 15.VI.1958; (Hm-2892), l 9, same locality and
collector, 21.VI.1958; (Hm-2703), l d: same locality and collector ,
25.VI.1958; (Hm-2751), 1 8; same locality, date and collector; (00002), 3
9, La slim, 1000 mn, Depto. del Valle, 6.II.l97ln, E. Méndez s L.
Velasquez; (00007), 1 6'2 9, Pichindé, Depto. del Valle, 21-25.l.l97u,
M. Thous and L. Velasquez; (00010), 1 9, same locality and collector,
7.111.1979; (00078); 3 6'9 9, Iago Celina, 1u50 m, Depto. del Valle,
22.111.1979, E. Méndez 5 L. Velasquez; (00079), 1 6'1 9, same locality,
date and collector; (00080), 6 0’5 9, same locality, date and collector;
(00080), 5 8‘5 9, same locality, date and collector; (00081), 1 9,

same locality and collector, 23.11.197u, (00082), 2 8'5 9, sane locality,
date and collector; (00083), 2 cm 9, sane locality, date and collector;
(00085), 3 do 9, same locality, date and collectOr; (00085), 3 o'5 9,
same locality, date and collector; (00088), In c”3 9, sane locality, date
and collector; (00089), 1 0'2 9, same locality, date and collector;
(00090), 3 6’1 9, sane locality, date and collector; (00101), 1 9,

samne locality, date and collector; (00102), 1 o”2 9, same locality,

date and collector; (MEI‘ ll265),01 o’,’ Florida, 8 km S.E. "La Diana,"

61

1700 m, Depto. de1Valle, 5.XII.1970, L. Velasquez; (MET 0257), l 9, sane
locality, date and collector; ex 'Ihonasomys fuscatus, (HIE-1672),

 

1 9, Pichindé, Mnmicipio de Cali, Depto. del Valle, 9.IV.1968, H. Trapido.
Rerarks. Our records of _P. bohlsi bohlsi extend the range
of this taxon which is row lcnom from Colombia, Ecuador, Venezuela,
Trinidad, Brazil, Argentina and Paraguay. The majority of our
southwestern Colombia specimens are from cryzorine rodents obtained from
1000 to 2000 meters elevation. Tipton alnd Machado-Allison (1972) suggest
that the optimum habitat of this flea is at elevations between 1000 and
1500 meters and the preferred hosts are cricet'me rodents and perhaps
more specifically of akodont stod< .

Polygenis caucensis , n. sp.

 

(Figs . 21, 22)

Emmaterial. Hloltype o from m caliginosus, (00036),
Alto Anchicaya, 550 m, Depto. del Valle, Colonbia, ll.II.l970, E. Méndez
8 L. Velésqnez; aJJotype 9, (00032), same locality, date and collector;
1 9 paratype, (00002), same locality, date and collector; 1 o paratype
(Aoc. No. B—57l), from Didelflg's marsgialis, Curiche River, Depto. del
Choco, Colonbia, 19.VI.1967, D. 5. Young. Holotype and allotype will be
deposited at the National Men of Natural History, Washington, D. C.;
1 paratype will be deposited in the British Mnoeum (Natural History) and
1 paretype in the Gorgas Melorial Laboratory's collection.

Dim. This speciesappearstobeclosesttofglygflroberti
;_be_e_be1;_'_ I. Fox, from which it may be differentiated by the more reticulate

 

In

I:

62

aedeagal lateral lobes. In addition, in P. caucensis, n. sp. fine
aedeagal median cbrsal lobe possess a subapical ridge which appears
tobeabsenting. robertibeebei.

DESCRIPTION.

MLE. _Hgfl (Fig. 21 A): Frons fairly rounded, interrupted by
short, angular tubercle protruded upward. Preantennal region with
moderate umber of micropcres and ‘2 pits in front of lst antennal
segment. Preocular row of 2 bristles inserted near eye. Post-ocular
row represented by few minute bristles located on lower and upper
portions of preocular region. Arch of tentcriun not conspicuous.
Oral angle well defined. Genal lobe seniangular. Eyes subovate,
not incised, large and well pigmented. Maxilla with acuminate tip
reaching last segment of maxillary palp. Post-antennal region with
3 rows of bristles behind lst antenrnal segnent. Antennae densely
covered wifin minute and prominent bristles irregularly distributed.

m: Pronotun and mesonotum wifin 2 rots of bristles.
Mesepisternun wifin 2 large bristles per side . Mesepimeron apparently
with 2 or 3 bristles perside. Metanotunhaving 3 rows of bristles.
Metanotal flange with about 6 spinelets . Metepisternun with single
bristle. Metepimneron with 9 bristles arranged in 2 rows .

BE: Posteromarginal notches of metatibia having strong
bristles with following distribution: 2-2-2-3—2-3.

Abchnen: Terga I-V with spinelets and 2 rows of bristles.

[humidified sterna wifin single row of bristles , those of sternna II and

:1)

63

III preceded by very few marginal and submarginal bristles.

Modified abdominal segments (Fig. 21 C):'5t VIII somewhat
reduced, provided with several unequal bristles. Fixed process of
clasper (Fig. 21 B) broad, but its total length exceeds maximum
width. This structure shows barely projected apical lobe, cbrsal
margin shallowly sinuate , posterior and ventral margins strongly
sinuate , fine last one moderately indented. Chaetotaxy as
illustrated. Movable finger of clasper (Fig. 21 B) not reaching
apical lobe, provided with several marginal and inconspicuous
imner bristles. Proximal arm of sternum IX shorter than distal arm,
of irregular shape, narrower at basal portion, finen becoming
considerable expanded and sinuate, terminating in subangular
projection. Distal arm of sternum IX (Fig. 22 A) curved cephalad,
more dilated medially but gradually tapering towards its subrounded
tip. Chaetotaxy of finis arm limited to some apical bristles and
postero—marginal bristles of various sizes. Heel of sternum 1X
prolil'ent, with slender, straight terminal tendon. Sternum VIII
wifin wide subrounded caudal lobe and single row of unequal bristles .
Apodeme of aedeagus (Fig. 22 B) broad, shorter finan terminal portion
of aedeagno, with subrounded apex. Terminal portion of aedeagus
(Fig. 22 B) With semiromnded distolateral lobes. Median heal lobe
large , almost reaching posterior portion of distolateral lobes , with
noper margin shallowly convex. Crochet not enparent. Lateral lobe
gently reticulated , broadly curved, except for short basal angular
prominence. Side piece prominent, almost triangular, its longest side
fairly convex. Fender well sclerotized, arched and conspicuous.

61+

Apical portion of inner tube larger than basal portion, strongly
convoluted, wifin final portion slender. Basal portion of inner tube
of moderate proportions, with narrow foreren on anterior half.
Crescent sclerite short, indistinct. Side piece broad, of irregular
shape, having anterior margin convex, posterior margin angulate.
Moral latero-ventral lobe represented by short, slightly curved
knob-like structure. Pseudo tube long, sinuate, well sclerotized.
Lateral finickening of end dnamber very long, sinuous, reaching heel
at base of aedeagal poudn. Heel conspiculus, slightly curved Inward,
wifin apex subrounded. Fluted membrane prominent.

FEMALE. Head, thorax, legs and unmodified abdmninal somites
essentially as in male.

Modified abchnrinal segments: Tergum VII ventrally expended
beyond longitudinal axis of abdomen, having 2 rows of bristles.
Tergun VIII large, bearing sinuous posterior margin and bristles of
different lengfin. Sternum VII with posterior margin subtruncate ,
provided wifin several uneven bristles apparently forming single roz.
Sternum VII broad but not conspicuous, provided with subangular caudal
margin. Dorsal anal lobe and ventral anal lobe both with subtruncate
apex and bearing several bristles. Anal stylet about two times as
long as maximum widfin, having 2 minute ventral bristles preceding long
apical bristle. Spermfineca (Fig. 22 C) wifin hulped bulga well
(blimited from short, upturned hilla. Dorsal margin of bulge and apex
of hilla respectively provided with short projection.

M: Holotype, 2.95 mm; allotype, 3.31 mm.

65

Remarks. The trivial name of this taxon has been adopted from
fine Depto. del Cauca where part of the type material was obtained.

Polygenis delpontei , n. sp.

Type material. Holctype o’and allotype 9 (Hm—315) gym
caligincsus, Colombia, Depto. del Valle, Municipio de Cali, Quebrada
Hmda near Pichindé, elevation 1800 m, 7.X.1967, H. Trapicb. Paratypes:
3 99 with same data as holotype; 1 d‘ (Hm-105) with same host, locality
and collector as holotype but 13.VIII.1965; l o’, l 9 (Hm-212) with
same data as holotype but 8.IX.1965; l 9 (Hm-213) from Reitlnro—
m mnexicanus, other data as HTC-212; l o’GflC-2u3) with same data
as HI'C-212 but 15.lx.1955; l 9 (Hm-302) from m (Oligoryzanys)

 

sp. ('2) with same data as HIE-212 but l(.X1965. Following paratypes
with same data as holotype except date: 2 carom-317) 11.x.1953;

1 9 (mo-330) 19.x.1955; l 9 (Hm-393) l7.XI.1965; 2 oo (Hm-397)
18.x1.1955; l Juno—029) with same data as holotype but 29.x1.1955;

1 9 (Hm-050) with same data as holotype but l0.-XII.l955; 1 6"(HI‘C-1307)
with same data as holotype but 13.XII.1966, 2 c’(00007) same host,
Rincan del Yarunal, Pichindé, Depto. del Valle, 21-25.I.l970, M. Thomas
5 L. Velasquez; l c'(00010), La Buitrera, Depto. del Valle, 7.11.1970,
E. lendez 8 L. Velésquez; ex W fuscatus, l d; (HIE—2955),
Rincon del Yarmal, Pidnindé, Depto. del Valle, 8.V.1969, H. Trapido,
l d; with same host and locality but 28.V.1959; 1 9 (00050), Saladito
(Km 12), 2000 m, Depto. del Valle, 17.11.1970, B. Méndez 5 L. Velasquez;

66

1 d' (HI‘C-805) with same data as holotype but elevation 1700-1900 m,
5.II.1966; 1 0"(HI‘C-1317) with same data as HI'C—BOS but 13.XII.1966.

Holotype and allotype will be deposited in the U. S. National
Museum of Natural History. Paratypes will be distributed among the
following institutions and specialists; British Museum (Natural
History), Universidad del Valle, Colombia, Gorgas Memorial
Laboratory Dr. Robert 'Iraub and Dr. Phyllis T. Johnson.

Dimis. Polygenis delpontei, n. sp. is closely related to
E. bradfinm Jordan. It differs fran this species primarily in that
it has the terminal portion of the aedeagal sclerotized inner tube
rob—like and upturned. In 1:. brachinus the termiral portion of the
sclerotized inner tube is flat and bent dowmard.

DESCRIPTION.

MALE. _He_a_d (Fig. 23 A): Prons evenly rounded. Frontal
tubercle barely projected out of margin, subrounded by wide
sclerotized area. Preantennal region profusely covered with
mdcropores. Preantenral row consisting of 7 unequal bristles evenly
spaced. Preoculm' row formed by 3 medium size bristles and 2 or 3
secondary short bristles. Premarginal bristles of antennal fossa
weak, inconspicuous . Oral angle acute but not prominent. Genal lobe
subanglflar. Eye oval, with small ventral indentation and moderate
pignentation. Post antennal regim essentially with 3 rows but
posteriormost bristle salamat displaced, thus suggesting |+th row.

Thorax: Pralotunandmesonottmneachbearing2rws of

 

bristles, last row with intercalaries . Mesepisternum SGTBWha't

67

reticulated, with 2 bristles. Mesepimerm with 3 bristles. Metanotum
with It rows of bristles; anterior row reduced to about 3 or 0 bristles.
Flange of metanotum having 9 to 2L1 spinelets .

_ngg. Typical of the gems . Metatibia with posteranarginal
notches bearing strong bristles arranged from upper to lower as follows:
2-2-2—3-2-3.

Modified abdominal segments (Fig. 23 C). Terglm VIII small,
having single row of bristles . Sternum VIII relatively large ,
caudally forms subangular lobe, with 1 row of 5 to 7 mequal bristles.
Caudal flaps of sternum VIII opening at 1/3 of ventral margin. Dorsal
and anal lobes of proctiger as in other Polygenis. Fixed process of
clasper (Fig. 23 B) with subangular apical lobe. Dorsal margin
barely sinuate , ventral and posterior margins strmgly simate but
lacking true indentations. Dorsal area of fixed process very setose;
rest of bristles of this structure scantly distributed over ventral,
posterior and inner areas. Movable process of clasper (Fig. 23 B)
slightly slorter than distal arm, showing broad apex, usually of 3
lobes. Distal arm of sternum D( (Fig. 23 A) curved cephalad, with
broadflbase, but having;about equalfiwidth throughoutimost of its length.
Its numerous bristles oriented posteriorly. Heel of sternum IX small ,
bearing short slender tenfiral tencbn. Apodane of aedeagus
(Fig. 20 B) with very broad portion before its subromded apex.
Terminal pcrtim of aedeagus (Fig. 20 B) having prominent rounded
distolateral lobes . Median dorsal lobe with posterior semiangular
projection. Crochet indistinct. Lateral lobe broadly convex. Dorsal
area of terminal porticn of aedeagus provided with chistinct striated

68

section facing apical portion of sclerotized inner tube . Lateral
thickenning of end chamber sinuous, ending almost at level with basal
segment of sclerotized inner tube. Fender long, sinuous and slender.
Basal portion of sclerotized inner tube much shorter than apical
portion of this tube, having large foreman on posterior half. Ribs
scant, with irregular distribution. Crescent sclerite short, sinuous .
Fulcral latero—ventral lobe curved caudad, with rounded annex. Heel

at base of aedeagal pouch subacute, connected with apodenal rod.
Vesicle ventrally expanded into thick ridge of irregular shape . Final
section of this ridge apparently receiving anterior portion of fluted
malbrane. Penis rod coiled, ending on thick nob-like portion. Fluted
mnenbrane moderately develOped.

W. _H_ea_;_<_l_ (Fig. 25 A): With frons more evenly rounded than
in male. Thoracic structures and legs similar to those of male.
TergnmeichorBrwsofbristles andpostermarginalrowof
spinelets. Other modified abdominal somites as in male.

Modified abdominal segments (Fig. 25 B): Sternum VIII
moderately developed, extended beyond longitudinal axis of abdomen,
having 2 rows of bristles and single antepygidial bristles . Tergun
VIII long, with posterior margin somewhat engular, provided with
several postero-marginal and submarginal bristles preceded by long
irregular row of bristles. Sternum VII with postero—caudal margin
abruptly truncate, not incised, raving a combination of long and short
bristles . Sternunn VIII reduced, caudally truncate . Sternun IX short,
with posterior margin straight or barely arched. Dorsal anal lobe of
proctiger (Fig. 25 C) with truncate apex and prominent apical and

69

subapical bristles . Anal stylet with 2 minute ventral bristles before
long apical bristle. Spermatheca (Figs. 25 B, D) having cribose
bulga perfectly separated fromn short hilla with terminal portion
Wd, dilated, semi-globular.

m. Holotype, 2.0!: mm; allotype, 2.01 mm.

Remarks. This species is dedicated to the memory of the late
Dr. Eduardo Del Ponte, whose excellent work on Siphonaptera and other
blood-sudcing insects contributed to the foundation of medical
entonology in South America.

Polygenis dunni (Jordan 8 Rothschild)
(Figure 26)

Rhopalcpsyllus diva; Jordan a Rothschild, 1922, Bctop., 1:209,
figs. 251, 262.

Remarks. Althoughthis specieshasrotbeenyetrecordedfrom
Colonbia, there is a strong possibility that it occurs in this country.
It is presently known from Panama, Venezuela and Trinidad, and
paresitizes an assemblage of hosts (see Tipton and PEndez, 1967,
and Tipton and Machacb—Allison, 1972).

Polygenis kinsi, n. sp.
(Figs. 27, 28)

RE material: Holotype male ex Ormys albigularis
(Hm-1838), Cerro Munchique (60 Joe by road west of Popayan); Pena
del Cerro, elevation 2160 m, Departamento del Cauca, Colonbia,

70

ll.V.1967, H. Trapido.

Holctype will be deposited in the U. 3. National Musenmn of
Natural History.

m: Polygenis hcplcninsi, n. sp. is similar to E.
M Jordan 8. Rothschild, from which it is readily differentiated
in having the terminal portion of the sclerotized inner tube bent
upward. In E. _l_i_t_a_rg_us_ this struotme ends barely sinuate and
oriented cephalad.

IESCRIPI'ION .

MALE. Head (Fig. 27 A): Prons rounded, its contour not
interrupted by clypeal tubercle which is not pronounced.
Preantennal area above clypeal tubercle covered with micropores.
Preantennal row of bristles consist of 5 bristles, the one near
antenna exceeds others in size. Preoculam row of bristles
represented by 3 principal. long bristles and short secondary bristles
separated from displaced minute bristle inserted near ventral margin.
Eye subovate , moderately pigmented, with deep ventral indentation .
Tentoriun with arched anterior arm preceding principal stem. Oral
angle short, acuninate. Tip of maxilla reading anterior portion of
last segment of maxillary palp. Genal angle subacuninate. Post-
antennal region having 3 rows of gradually increased number of bristles.
Micropores limited to space between antennal base and first row of
bristles. A group of short and thin bristles located near scape of

antenna.

71

lh_gr__ax_: Mesonctumn bearing 3 rows of bristles, first row reduced,
containing 2 or 3 bristles. Mesepisternum with 2 bristles.
Mesepimeron with 3 bristles . Lateral metanotal area having 2 large
bristles preceded by 2 short bristles. Metepisternum with single
long bristle and group of short bristles on anteromarginal projection.

legs: As in other Polygenis . Metatibia with strong bristles
inserted in notches as follows: 2-2-2-3—2-3.

Abdomn: Tar-gun I with 3 rows of bristles, lst row reduced
to about '4 bristles. Renaining unmodified terga with 2 rows of
bristles. Sternum I with scattered ventral, subventral and
inner bristles. Other unmodified sterna with single row of bristles.

lbdified abdominal segments (Fig. 27 C): Tergum VIII short but
reaching abcbm'nal axis. Sternum VIII large, having truncate
caudonarginal expansion, its ventral division well posterior to row
of bristles. Fixed process of clasper (Fig. 27 B) showing dorsal
marg'n moderately sinuated, gradually elevated towards apical portion .
Posterior margin of fixed process very undulate, having slort semi-
angular protrusion proudmad to fovea of fixed process. Ventral margin
deeply indented. Bristles of fixed process as illnatrated. Movable
finger (Fig. 27 B) slightly bent upwards, not reading apex of fixed
process of clasper, clothed with several marginal, submarginal and
inner bristles. Distal arm of sternum D( (Fig. 28 A) semi-falcate,
almostas longas proximal arm, lnaving anteriormarg'n devoidof
bristles and posterior margin with bristles of moderate size
distributed over half its length. Heel of sternum 1X heavily
sclerotized, bearing relatively short tendon. Distolateral lobes of

72

aedeagus (Fig. 28 B) with anterior portion subrounded and posterior
portion obtuse, prolonged into large ventrcmarginal extension. Median
dorsal lobe having subangular apical portion. Apicomedian sclerite of
aedeagus striated. Crochet imperceptible. Lateral lobe undulate ,
showing distinct superior subangular projection. Dorsal area of
terminal portion of aedeagus with small striated section facing
junction of apical and basal portions of sclerotized inner tube .
lateral thickening of end chamber sinuous, ending at level of vesicle.
Lateroventral sclerite sickle-shaped but with rounded tip. Heel at
base of aedeagal pouch semi-acuminate , attached to apodemal rod.
Fender semi-arched, slender and elongate . Apical portion of
sclerotized inner tube with 2 100ps, its terminal section wide,
upturned, oriented cephalad, provided with semiglobular apex. Basal
portion of sclerotized inner tube having mesal foreman of moderate
size. Ventral nodular section of basal portion large and prominent.
Ribs distributed from an area near vesicle to fender. Crescent
sclerite arched, not prominent. Side pieces semi-triangular, with
opposite ends acuninate .

m: Holotype, 2.31 mm.

Remarks. This species is named for the late G. H. E. Hopkins
in recognition of the outstanding contributions he made to the
systenatics of Siphonaptera and Anoplura .

Polygenis klagesi (Rothschild)

 

(Figure 29)

73

Max klagesi Rothschild, 1900, Novit. Zool., 11:620., Pl. 9,
fig. 28; P1. 10, figs. 35, 39.

Material examined: Ex Proechimys semispinosus, l 6', Rio
Raposo, Depto. del Valle, XI.1959, P. Barreto; ex Hoplomys grumurus,

 

(00016), 5 6'5 9, Alto Anchicaya, 550 m, Depto. del Valle, 10.11.1970,
B. I‘findez 8 L. Velc’nsquez; (00017), 2 0’3 9, same locality, date and
collectors; (00018), 3 o"1 9, same locality, date and collectors;
(00019) 1 6", same locality, date and collectors.

In addition to the material from the southwest part of Colonbia,
I have emnined 11 males and 17 females from Carinagua, Depto. del
Meta, 8 males and 11 females from fine Depto. de Antioquia and large
series of males and females from Depto. del ro6.

Remarks. Our specimens of Polygenis klagesi taken on
M murus from Alto Anchicayé, Depto. del Valle, and Curiche,
Depto. del Ctooi, do rot agree in certain features wifin specimens
taken on Proechimys senispinosus which we are tentatively

interpreting as typical Polygenis klagesi samnuelis. In View of the

 

considerable amount of variation displayed by 1:. @533, it is
advisable not to try to segregate Colombian populations into
subspecies until an adequate study of material obtained finroughout the
geographical range of these fleas is done.

Polygenis klagesi is presently known to occur in Brazil, Panama',
Costa Rica, Colombia, Venezuela, Trinidad and Ecuador, from sea level
to altitudes below 900 meters.

'Ihefi. ]d_ageiicomplexoccurs onan spectumofl'osts; however,

these fleas are more naturally associated with the spiny rats family

70

Echimyidae, particularly with Ptoechimys.

Polygenis pgadoi (Wagner)

 

(Figure 30)

Rl'opalopsyllus predoi Wagner, 1937, Zeits. Parasit., 9:020,
fig. '4.
Material examined: Ex Oryzonys albigularis, (RFC-2780),

 

1 (5‘2 9, Pichindé (La Esperanza), 1900 m, Municipio de Cali, Depto.

del Valle, 20.v111.1959, H. ‘I‘rapido; (Hm-2186), l d", Pichindé, 1900 m,
Municipio de Cali, Depto. del Valle, 25.1.1968, H. Trapido;

ex Oryzonys caliginosus, (RIC-289), l 9, Quebrada Honda near Pichindé,
1000 m, Municipio de Cali, Depto. del Valle, 30.D(.1965, H. Trapido,
(Hm-310), 1 9, same locality and collector, 5.x.19ss; (Hm-315), 1 9,
same locality and collector, 7.x.1965; (Hm-707), 1 o"0 9, Valle del
Rio Pichindé, 1700-1900 m, Municipio de Cali, Depto. del Valle,
28.VI.1966, H. 'hrapido; (Hm-005), 1 9, samne locality and collector,
s.v11.1900; (Hm-700), l 0", same locality and collector, 16.VI.1966;
(mo-032), 1 9, Pichindé, 1800—1880 m, Municipio de Cali, Depto. del
Valle, VII.1966, H. Trapido; amt-902), l 9, Valle del Rio Pichindé,
1700-1900 m, Depto. del Valle, H. Trapido; (Hm-075), 1 d3 9,
Pichihdé, 1000-1090 m, Municipio de Cali, Depto. del Valle, 7.VIII.1966,
H. Trapicb; (Hm-090), 2 9, same locality and collector, VIII.1966,

H. Trapido; (Inc-902), 1 62 9, Valle del Rio Pichindé, 1700-1900 m,
Municipio de Cali, Depto. del Valle, 22.VIII.l966, H. Trapido;
(two-1000), 1 o”0 9, Valle del Rio Pichindé, 1700-1900 :11, Municipio

75

de Cali, Depto. del Valle, 13.1x.1955; (Inc-1001), l 9, Pichindé,
1780 m, Mmicipio de Cali, Depto. (bl Valle, 1H.X.1966, H. Trapido;
(Hm-1109), l 9, Valle del Rio Pichindé, 1700-1900, Mnnnicipio de
Cali, Depto. del Valle, 20.x.1955, H. Trapido; (RIC-1262), l 6’; same
locality and collector, 10.x1.l955; (Hm-1250), l a: same locality
and collector, 2l.XI.1966; (HTC-l283), l 0’; same locality end
collector, 5.XL1966; (RFC-1283), l 6; same locality and collector,
5.XI.l966, H. Trapido, (mo-1307), 2 9, same locality and collector,
13.x1.l955; (Hm-1002), 1 0’1 9, same locality end collector,
20.1.1957; (Hm-1090), l 9, same locality and collector, 15.111.1957;
3 d0 9, Pichindé, 1900, Municipio de Cali, Depto. del Valle,
VI.197'+; ex Rhipicbmys latimanus, (Hm-1229), l c"; Pichindé, 1900 m,
Muncipio de Cali, Depto. del Valle, 17.x1.l955; H. Trapido ex
Thomasogys fuscatus, (Hm-1220), l 9, Valle del Rio Pichindé, 1700-
1900 m, Municipio de Cali, Depto. del Valle, 7.XI.1966, H. Trapido.

Remarks. Polygenis mpg; has been reported before from
Brazil. Our specimens from southwestern Colonbia represent fine first
records of finis flea for fine country. Orstonys (Melanoma)
caligfi us, probably the cormonest rodent in southwestern Colonbia,
stands outasfinemore favoredhostinfinis territory. Of 08
specimens of g. piagpi obtained, I42 were from finat host while four
were takenon m albigfl's, oneoanatimams
endoneon'I'tomasomfifuscatus. Thereportsoffinis speciesfrom
Brazil concern the following l'osts: Eng; socialis, Didel 's
cancrivora, mmwes, Aho___gc_rn_ sp. (possibly cu___s_g_r_'), a wild
rat and wild mouse (Johnson, 1957).

76

Polygenis roberti beebei (1. Fox)
(Figure 31)

Rhopalolngwllus beebei I. Fox, 1907, 2001: N. Y. 2001. Soc.,
32:117, fig. 2.

Remarks. Polygenis roberti beebei has not been yet collected
in fine soufimwest corner of Colombia; in spite of finis fact we consider
its presence finere as probable. This taxon has not been reported from
Colonbia in the literature; however, we have examined 36 specimens
(15 males and 21 females) from the Departamento de Antioquia,
kindly loaned by Dr. v. J. Tipton.

The zoogeography of _R. roberti beebei presently involves
Venezuela, Per'fi, Colombia, Trinidad and Panama. It has been collected
end low end moderate alt'tudes. The normal Tosts of this flea are
marsupials and rodents, notably sore members of the cricetine

gems 922m-

Polygenis thurmnani Johnson

(Figure 32)

Polyggn_is finurmani Johnson, 1957, Men. Ent. Soc. Wash. 5:169-
170, P18. 81%, 85.

Material examined: Bx Didelphis marsupialis, (00093), l 62 9,
Iago Calima, 1950 m, Depto. del Valle, Colonbia, 23.11.19714, E. Méndez
8 Velasquez; ex MM (RFC-2769), l 0’; Quebrada Ncrte,
Finoa La Flora, 1900 m, Pichindé, Depto. del Valle, 1.VIII.1968,
H. 'I‘Iapido; (HIE-2779), l o’; la Esperanza, Pichindé, Depto. del Valle,

77

20.VIII.1968, H. Trapido; (HTC-160), l 9, Quebrada Honda near
Pichindé, 1500 m, Municipio de Cali, Depto. del Valle, 17.VIII.1965,
H. Trapido; (HI‘C-301), 1 9, same locality, date and collector,
l&.X.l965; (RFC-1159), l d', sere locality, date and collector,
15.x11.l955; (Hm-1000), 1 9, Valle del Rio Pichindé, 1700-1900 m,
Municipio de Cali, Depto. del Valle, 0.x.l955; (00057), 3 d1 9,
Pichindé, 1600 m, Depto. del Valle, 21—25.I.197|+, M. Thomas 8 L.
Velésquez; ex m sp. (probably 9. caliginosus), l 9, Quebrada
Honda near Pichindé, 1800 m, Municipio de Cali, Depto. del Valle,
10.XII.1955, H. Trapido; ex Rhipidomys latilmanus, (HTC-BlS), l c',
Valle del Rio Pichindé, 1700-1900 m, Municipio de Cali, Depto. del
Valle, 12.VII.1966, H. Trapido; (Hm-2550), 1 9, Finca La Maria,
1500 m, La Cumbre, Depto. del Valle, 1.XI.1967, H. Trapido.

Rerarks. Polygenis thurmani is no: known from Perfi end
Colombia. The vertical distribution indicated by our material from
southwestern Colombia fluctuates from 1050 meters to 1900 meters
elevation. Our present information shows that in every one of these
countries fine hosts favored by g. thumani are different. In PerCn
this flea has been taken from Akodon pulcherrimus inambari , Phyllotis
m, 9m stolzmanni or O_xym_vcteris p_. nigifrons. Our
Colombian specimens are from Didelphis marsupialis, Oryzomys
albigularis, Oryzomys sp. (probably _0_. caliginosus), and Rhipidomys
latimanus. The data on finis species is still inadequate for drawing
any conclusions on host preference . Pbdever, the premise that
m albigularis is fine typical lost might be reasonably founded

78

on fine basis of our collection. Ten out of 16 specimens of B.

finurmani. were found on this rodent.

Polygenis trapidoi, n. sp.
(Figs. 33, 30)

mmaterial. Holotype o (RIC-1236) from Oryzogrys caliginosus,
Czflombia, Depto. del Valle, Municipio de Cali, Valle del Rio Pichindé,
elevation 1700-1900 mu 31.X.1966, H; Trapido. Paratypes as follows:

1 o’with same data as holotype: l d’(HTC-1629) with other data as
holotype but 31.III.1967, l d' (RIC-1603), represented by mounted
genitalia only, with same data as holotype but l7.III.1967; l c”
(00005), same host and locality but 21-25.197|+, M. Thomas 8 L.
Velasquez; l d’(00007), same data as 00005, 1 c’(00005), same data
as (00005).

Holotype will be deposited in fine U. S. National MUseum of
Natural History. One paratype will be deposited in the BritiSh
iMbseumn(Natunal History) and the othernparatypes will remain in the
Gorgas Memorial Laboratory's collection.

M3. Nearfi. gu_nn_i_Jordan 8 Rothschild fromwhich it
is separable by the subangularnonkaauginal expansion of the sternum
VIII. Other diagnostic structures are contained.in.the genitalia

illustrated.

DESCRIPTION.

79

MALE. Ileag (Fig. 33 A): Frontal tubercle not exserted.
Micropores dispersed from area before falx to prondmity of frontal
tubercle. Preantemal row of bristles consisting of 5 bristles,
the one near antenna the longest. Preocular row having 3 prominent
bristles and about equal number of minute bristles. Oral angle
acnminate . Bye nodemtely pigmented, showing profound ventral sinus .
Tentcriun well defined, having anterior sten strongly arched.
Maxillary lobe acuninate, reaching lnth segment of umdllary palp. Genal
lobe definitely angular. Post antennal area with 3 rows of bristles
and group of minute bristles close to antennal scape. Pronotum wide,
its posterior margin reaching beyond level of proepimeron .

M: Pronotnnn and mescnotum each with 2 rows of bristles.
Metanotun having 3 rows of bristles . Lateral metanotal area
possessing 3 short anterior bristles followed by 2 large posterior
bristles . Metepisternum with single bristle near pleural arch.
Metepineron with 6 bristles distributed in 2 rows.

135E: Metatibia with 6 posterunarginal notches bearing strong
bristles as follows: 2-2-2-3—2-3.

Modified abdominal semts (Fig. 33 C); Tergum VIII small
but reaching abdoninal ast. Sternun VIII slaving posterior margin
undnflant but entire , with appreciably expanded subangular
pcsterocaudal portion, having 10 bristles of diverse sizes oriented
on mesal area. Fixed process of clasper (fig. 33 B) provided with
subangular posterolateel elevation. Posterior margin of fixed
process with angular prominence at about l/ll of distance from top.
Ventral nargin notably undulate disu'ibution of bristles as illustrated.

80

Movable process of clasper (Fig. 33 B) elongated but not reaching
apical portion of fixed process of clasper; barely wider on medial
region with lateral nargins slightly sinuate , having bristles
scattered along margins and inner areas. Prondnnal arm of sternum IX
(Fig. 33 C) shorter than distal arm, constricted before bilcbed apex.
Distal arm of sternun IX (Fig. 30 A) elongated, slightly curved
upward, with numerons marginal and submarginal bristles of various
sizes. Basal Spur of sternum IX projected, showing thick basal
portion and highly sclerotized, provided with slender tendon. Disto—
lateral lobes of ternfinal portion of aedeagus (Fig. 30 B) somewhat
falcate. Median dorsal lobe angular. Apico—median sclerite of
aedeagus reduced and striated. Crochet indistinct . Lateral lobe
wide, irregularly sinuated. Fender reduced and arched. Apical
portion of inner tube slightly larger than basal portion of sclerotized
inner tube, with coiled section of 2 or 3 loops, ending in free
undulated portion directed mud. Basal portion of sclerotized inner
tube compact, with most of superonarginal area straight, having very
narrow foreman almost limited to anterior half but extended beyond mid
point into fraction of posterior half. Ventral nodular section of
this basal portion small. Ribs very nunerous, distributed on most of
anterior section of terminal portion of aedeagus . Crescent sclerite
sinuated, inconspicuom . Side pieces elongated, sinuous , having 2
opposite acuminate extremes . Latero-ventral sclerite represented

by curved knob-like snucture with broad cbtnse apex. Phel at base of
aedeagal pouch send falcate, very prominent. Fluted membrane well
developed.

81

Egg. Holotype, 2,0“ nun.

Remarks. I take great pleasure in dedicating this species to
Dr. Harold Trapido in appreciation for his kind collaboration and his
contributions to the knowledge of ectoparasites .

Scolopsyllus colonbianus Mendez

 

(Figs. 35, 36)

Scolmsyllus colonbianus Méndez, 1968, J. Med. Ent. 5:005-010,

 

Figs. l-llt.

Material examined: The following data include the type material.
Ex m caliginosus, (ETC-207), o'holotype and 9 allotype, Quebreda
Honda near Pichindé, Municipio de Cali, Depto. del Valle, 1800 m,
0.13:.1965, (mt-051), 1 9 (paratype), same locality and collector,
l3.XII.1965; (Inc-619), 1 9 (paratype, same Departamento and Municipio,
but from Valle del Rio Pichindé, 1700-1900, 5.1V.1966, H. Trapido;
(Int-1076), 1 9 (paratype), same locality and collector, u.III.1957;
ex 952% alfaroi, CHIC-126D, l o" (panatype), same locality and
collector, 1+.XI.1966; ex Didelphis marsupialis, (71-625), 1 9,
Pichindé, 1600 m, Depto. del Valle, 6.VIII.1971, H. Tyndale-Biscoe.

. Remarks . Scolopsyllus colcmbianus is probably endenic to sub-
tropical nountains of moderate elevations surrounding the Cauca Valley.
The preferred host appears to be Oryzorys caliginosus, which is the most
abundant and widespread rodent in the Departanentc del Valle.

Rhcpalopsyllus australis tupinus Jordan 8 Rothschild

82
(Figure 37)

Wmuus australis tupime Jordan 8 Rothschild, 1923,
Bctop., 1:328, fig. 339.

Material examined. Bx Eire barbara, l o", Pichindé, Depto. del
Valle, 10.VI.197'-l, M. D. Little.

Renarks. Data gathered in Panama (Tipton and Méndez, 1966)
indicated finat _R_. australis 1_:_up_inus is normally fornd in association
with the cavionorph rodents Dasyprocta punctata and m paga,
which are fine more selected host. The Collared Peccary, Tayassu
E3315, and fine White-lipped Peccary, Tgassu m, follow finose
rodents in host preference. Other anirrals such as Didelphis
narsupialis, Chironectes minimus, Pnoechinys senispinosus and other
rodents, are also favored to a lower degree. Occasional hosts are
carnivores like _Big barbarn, Galictis allamandi, _Cgr_ni_s_ familiaris,

 

and others, that are infested by fleas obtained fron their prey.

Rhopallcpsyllus australis minus has not been previously
reported fron Colonbia. We lnave also enamined one male and five
feneles fron Carimgm, Departanento del Meta. This flea has been
Imam before fron Panama, Bolivia, Brazil and PerCn. At least in
Panama, this taxon has been collected fron sea level to elevations
close to 1600 meters.

Rhopalopsyllus cacicm saevus Jordan 8 Rothschild
(Figure 38)

83

Rnopalcps'yllus cacicus saevus, 1923, Ectcp., 1:325, fig. 332.

Remarks. Ihavenotbeenabletocbtainthis taxoninthe
sorfinwest part of Colonbia; however, it sees reasonable to presune
its presence in this territory since it is lonown from other areas
of Colonbia with similar ecological conditions and hosts. This form
has been reported fron finis country by Puller (19142) and I have
examined specimens from Carimagua, Departamento del Meta.

So far 3. cacicns saevus is known fron Venezuela, Trinidad,
Colonbia, Panama, Costa Rica, El Salvador, Guatenala and México. It
is apparent finat this flea has an affinity for Dasypns novemcinctus
and Didelphis mfialis throughout its geographical range;
nevertheless, it is interesting to note that our series from Carirragua
(12 males and 5 females) were entirely obtained from Agonrti paca. In
Panama Jurtapulex ednidncphagcides seers to replace 3. cacicus saevns
on its principal host, D_a_syp_n§ novencinctus fenestratus in areas of
high elevations (Tipton and Mendez, 1967).

Rlnopalqpsyllus 1_ug_\_1b__r__is Jordan 8 Rofischild
(Figure 39)

Rhopalopsyllus _lpgu_bn_'_s Jordan 6 Rofinschild, 1908, Parasit.,
1:70, Pl. 3, fig. 12; Pl. 6', fig. 9.

Renarks. Alfinough specimens of R. Mare lacking in our
material fron fine southwest portion of Colonbia, finere is no doubt that
this flea occurs in this territory.

81$

Rhopalcpsyllus lugubris has been found in Panama, Colombia,
Brazil, Ford and Venezuela, parasitizing a miscellaneous group of hosts
at low and moderate elevations. The strong preference of this flea for
M Ec_a_ is shown in data presented by Tipton and Méndez (1967),
and Tipton and Machado—Allison (1972). Our collection of Colorbian
fleas contains a lot of R_. Mmpresented by 36 males and 70
females collected nainly from Agouti Raca at Carinagua, Departanento
del Meta. We are assigning these fleas to the form 5. lgggris
cryptoctenes (Enderlein).

SUPERPAMILY PULICOIDFA
FAMILY PULICIDQB
suammx PULICINAE
TRIBE momma
Xenopsylla Ms (Rothschild)

(Figure '40)

Pulex cheopis Rothschild, 1903, Ent. Mon. Mag., 39:35,
Pl. 1, figs. 3, 9; Pl. 2, figs. 12, 19.

Material examined. Ex Aie_le_s_ fusciceg, 9 6'12 2,
Buenaventuna, Depto. del Valle, l7.XI.1969, V. B. 'Ihatdner; ex
M calim‘ us, (ETC-862), l 9, Valle del Rio Pichindé,
1700-1900, 2.VIII.1966, H. Trapido; ex rat (probably m rm'vegicus),
(AA-29), 1 9, Vicinity of Cali, VII.196|+, A. Arata; ex mm,
(DEB 180 A), 1 d' 7 9, CIAT, Aoarology Barns, Depto. del Valle,
lu.XI.1975, J. B. Espincsa; (DEE 185 A), 3 0'5 9, sane locality,
XI-1975, J.R.E.

85

Renarks. Xenopsylla checpis is a comnon parasite of the
couuensal rats and is widely spread over the Asiatic, Efini0pian,
Palearctic, Neotropical and Australian Region. Many manuals have been
found parasitized by this major vector of plague, but murids occupy
first rank in host preference.

TRIBE ARCHAEDPSYILINI
Ctenocephalides felis (Douche)

(Figure l+1)

Pulex felis Boudné, 1835, Nova Acta Phys. Med. Acad. Caes.

 

Loop. Carol. 17:505, fig. 2.

Material examined. Ex Didelphis nmsupialis, (ETC-956), 2 d;
Vicinity of Cali, Municipio de Cali, Depto. del Valle, VIII.1966, H.
Trapido; (mt-2700), 2 d; Sonso, Municipio de Buga, Depto. del Valle,
25.VI.1968, H. Trapicb; ex gong @1318, (Hm-1010), 1 o", Universidad
Del Valle, Depto. del Valle, 25.V.1973, P. Barreto; (Inc-1060), 1 9,
sane locality and collector, 29.11.1971), ex M calijgnosus,
(00002), 1 9, Lago Calima, 1050 111, 23.11.1970, B. Mendez a L. Velasquez;
ex gigging, (DEE—186 A), 2 6'1 9, Depto. del Valle, 17 a 21.XII.
1975, D. E. Evans; ex Ming, (DE 183 A), l o”, Yulbo/
Palnesca Roadsise, Valle del Cauca, Depto. del Valle, 26.IX.1975, D. E.
Evens.

Remarks. This species is alnnost cosmopolitan andhas been
reported from sea level to over I4000 meters elevation. It has been
found on a wide range of hosts, including man; however, it is more
specific on fine donestic cat, §9_1i_§_ (Leg) Qty-i, and

86

secondarily on other carnivores .

TRIE PULICINI
Pulex irritans Iinnaeus

(Figure '42)

32.131 irritans Linnaeus, 1758, Sist. Nat., 10th ed.:61ln.

Renarks. Pulex irritans is cosmopolitan and inhabits warm and
temperate regions of the world. It has a wide host range and shows
marked preference for various carnivores, man and certain ungulates.
This species was reported from the Departamento del Valle by Renjifo
(1900). Other Colombian records are given by Puller (1902). gu_l_e_x_
irritans was considered the only flea (other finan 35.. che__op_is) of

inportance in Peruvian plague (Moll and O'Leary, 19u5:168).

Pulex simulans Baker

1113st Baker, 1895, Can. Eht., 27:65,67.

Material examined. Ex 'Ihonasolys cinereiventer, (HI'C—2u22),
1 9, Laguna de la Cooha, 2700 m, Depto. de Narifio, 18.V.1968, H.
Trapicb.

Fanatics. This specinen represents fine first record of
P. simulens in Colombia. The geographical distribution of this

species also includes Penami, Mécico, Venezuela and the United
States of America.

SUBFAMILY 'I‘UNGDlAE

87

Bhynchopsyllus pulex Haller

(Figure ll3)

Rtyndopsyllus pulex Heller, 1880, Ardn. Naturg., Jahrg.
06, Bd. 1:02, P1. 5, figs. 1-13.

Material examined: Ex Molcssus naior, (o-uas), 1 9, Site l-c,
Vicinity of Cali, Depto. del Valle, 8.VII.19611, A. Arata; (c—qu),
1 9, sane locality, date and collector; (c—980), 1 9, same locality,
date and collector, 23.VII.196ll; (c-982), l 9, sane locality, date
and collector; ex Molossus major lor Glossgphaga soricina lor Desmodns
rotmdus, 3 9, Site 1-A, Vicinity of Cali, Depto. del Valle, u.VIII.
1950, A. Arata; ex Noctilio labialis, (Pool N L 3), 3 9, Side of Rio

 

Cauca, Municipio de Cali, Depto. del Valle, III.197M, S. Ayala;
(NL-100F3), s 9, sanne locality, date and collector; (NL-looos-a), 6 9,
same locality, date and collector; ex Molcssus nolossun, ll 9,
Mayagnez, Depto. del Valle, v.197u, s. Ayala; (Hm-101), 2 9, (inside
house), Cali, Municipio de Cali, Depto. del Valle, 23.IV.1965, H.
'I‘rapicb; ex guano of Molcssus nnolossus E2122: 6 a", on attic of house
in "La Buitrera," approximately 10 km s. w. Cali (elev. i 1100 m),
Depto. del Valle, 19.IV.1975, M. Thales.

Remrks. The known geographical range of R_. pllegincludes
Argentina, Bolivia, Brazil, Chile, Colonbia, Bcuacbr, Peru, Venezuela,
Panama and Texas (southern United States). Inasmuch as this flea
is a true parasite of bats, especially nolossids, its distribution
is probably correlated with that of its hosts .

Dr. Maurice Thonas kindly presented to us six male specimens
of Wyllus that he reared fron guano of Molcssus molossus naicr

88

obtained at "La Buitrera," near Cali, Depto. del Valle. Comparisons
with males of Panamanian Rhynchopsyllus megastigmata, indicate that
they represent the same taxon. An evaluation of the characters
estimated as critical for separating fenales of R. p113; and R_.
Egastignata, allowed us to observe finat they are variable and
finerefcre unreliable. In view of these reasons, we conclude that
Rhyndnopsyllus megastigmata 'Iraub and Salmons, 1950, is a synonym of
Rhynchopsyllus pulex Haller, 1880 .

'I‘unga penetrans (lirnnaeus)
(Figure 00)

29135 pgnetrans Iinnaeus, 1750, Syst. Nat., 10th ed.:61|+.

Renarks. 103g; m is distributed in warm territories
of the Neotopical and Ethiopian regions . This species is adapted
to fixedparasitismandhas beenrecoveredfromvacious hosts but
display high preference for Sus scrofa and Hono sapiens .

 

Wehavebeenunabletoobtainspecinens ofthis fleafrom
southwestern Colonbia; however, the chigoe was reported from the
Departmento del Valle by Renjifo (1900). Dr. David E. Evans of
Centre Internacional de Agriculture Tropical, has donated to as six
specimens takenonSgMatFincaLolrdes, alocalityinthe
Departamento de Corcbba.

ANALYSIS OF inS‘I'ePARASI'I'E RHATIONSHIPS

Considerable information on fine host-parasite relationship
of Neotopical fleas is presented by Johnson (1957) and Wenzel and
Tipton (1966). Other publications dealing with finis subject are
I'blland (1961*), Pbpkins (1957), Pbpldns and Rofinschild (1953, 1956,
1962, 1966, 1971), Levis (1972, 1973, 1971* a, b, c, 1975), and
Tipton and Méndez (1966). Brief cements on fine flea taxa considered
hereinarechrivedfronthese andothersources, aswellasny
observations.

The available data concerning fine fleas of sorthwestern
Colonbia are not sufficient for an exhamtive analysis of zoo-
geography and host-peasite relationships . Additional collections
that contribute information on hosts , distribution and relative
abundance, will be necessary to answer the remaining questions
regarding fine ecology of finese parasites . Despite such disadvantages
sane general preliminary assmptions and conclusions can be presented
on patterns of dispersion and host-parasite associations .

In addition to natural host affiliation, tenperature, humidity
andofinerecolcgical factorsarecrucialindispersionoffleas
and other ectoparasites. Climate is probably one of fine most
inpcrtant factors that limits distribution of many species . Tenperate
areas arenotonlyricherinspecies cffleas finanthe lowlands,

89

90

but also have a great deal of endemism. (It may be observed in Table

2 that the majority of the flea taxa are concentrated on the Andean
territory). Of 35 taxa estimated for southwestern Colonbia, sons

12 species are characteristic of tropical zones (from sea level to

600 neters), while those remaining are typical of terperate or
subtropical zones (from 000 to 1,505 meters) and the much cooler
nontane areas (1,505 to at least 3,050 meters).1 Certain species found
inwarmareas are tolerant ofawide rangeintemperature, as
exemplified by Xenopsylla cheopis and ‘Ctenocephalides £e_li_s_.

Since fine evolution of fleas to some degree has paralleled
finat of their hosts, fine origin and dispersal of these nanmals can
often be indicative of fine distribution of their fleas. Pron the
argunents of some authors, rnotably Hershkovitz (1967) and Tipton and
Wenzel (1m, giL), it seens probable finat snall normals, particularly
cricetine rodents , dispersed recently (Miocene or earlier) from Middle
to Scufin Anerica. It is likely finat fineir fleas evolved and radiated
accordingly. Today, several genera of fleas, belonging nainly to the
families Ceratcphyllidae and Hystrichcpsyllidae , are associated with
finese rodents. In sane oases, adaptive radiation of Neotopical
rodents seen to have strongly influenced fine evolution of fineir fleas .
For ample, fine flea Kohlsia falcata Méndez and Hanssen, which occurs
in Colonbia, and l_<, M Mendez and Altan, a closely related
Central American species, have developed (along characteristics not

 

1
The altitudinal distribution need here follows Wenzel and Tipton

(1966).

91

shared with ofiner Kohlsia) a tibial ccub of spines on all legs. Since
fine two species are associated with sane manmals (squirrels arnd other
rodents) finat are either completely or partially arboreal, it seems
very likely that fineir peculiar adaptations are linked to features

of fineir hosts.

The genus Kohlsia is believed to have originated in Middle
America. The great majority of species of finis ceratophyllid group
occurs in Central America and Mexico, while only two species are
lancwn fran South Anerica (not further south than Colonbia and Ecuacbr).
This fact may indicate recent invasion of the gems to this part of
fine continent. A similar pattern of dispersal is found in the related
genus Pleochaetis. This group appears to have diverged from Middle
Anerica as typical parasites of fine sierle-penis—type cricetine
rodents (discussed by Hershkovitz, 1966), specifically Pergnyscus. It
is noted finat sane canplex-penis—type cricetines, such as M
and W, actually harbor fleas of fine genus Pleochaetis ,
possibly as secondary hosts. At fine present time, species of
Pleochaetis are distributed from soufi'mestern United States to
northern Soufin Auerica, but they are concentrated in Middle America.

The flea fauna of fine lowlands of Colonbia is very similar to
finoseofPanamSandfineAmazonandOr-inocobasins. (Fleaelenents
which reveal finis phenanencn are Acbrebopsylla (Trijtomylla)
internedia, Polygenis _k_l_.ag§_s_i, g. dn__r_ln_i_, g. roberti be__e_b_ei_,
Rhopalopsyllus auStralis, 3. cacicus and _R_. M) . This affinity
canbeunderstoodcnfinebasis ofcammspeciesorrelatedfcrus of
manual hosts, and fine absence of effective barriers finat might have

92

prevented faunal interchange since remote times . Conversely, the
strildng nnanmnalian relationship finat endsts between the Central and
Western Cordilleras in Colonbia and the Ecuadorian nountains is
correlated wifin the affinity found in the flea fauna. Species such as
Cleopsylla manticola , Sphinctogylla tolmerna, Plocopsylla phyllisae ,
Pleochaetis smiti, E. eguatoris and Tetrapsyllus comis , are exanples
of this similarity in the northern Andes.

In the soufimestern Oolcnbian fauna, coexistence of two or
more flea species on the same host is prevalent, and, in general,
host specificity is not highly developed. Phnever, exceptions are
Sternopsylla distincta sgciosa and mynchgpsyllus pulex, restricted
to bats. Even in such cases, not a single Chiroptera is the only
host for any of finese flea species. It is reasonable to assume finat
finese two bat fleas originated from fine northern half of South
Anerica, where fine hosts probably evolved.

0f fine cosnopolitan fleas fournd in southwestern Colonbia,
Xenopsylla dnfl's and Ctenocephalides felis are native to the
Ethiopian Region. LeptopsLlla aegis is derived from fine Oriental
Region (Hopkins and Rothschild, 1971), and finere is a strong
possibility finat ;P_n11_e_x_ irritans is derived from the Neotropical Reg'cn.
fu__1e_x_ sinnflans, which is restricted to America, as well as the other
species of fine genus, also could have fine same origin.

Ihsypsyllus gallinulae perpinnatus is South American, being
derived from the nanninate form which is probably fblarctic in origin.
It is logical that finis flea, like ofiner bird fleas, evolved on a
manual host (perhaps an arboreal member of fine family Cricetidae or

93

Sciuridae) before becaning adapted to birds .

Ofiner fleas, g. g. , Ctenidiosanus, which originated in South
America, might have evolved on cricetine rodents when they becene fully
established in finis portion of fine continent, or were early parasites
of cavianorphs or opossuns . This gems generates special interest
inviewoffine fact finat it is thecnly SouthAmericen groupofthe
more typical Old World family Pygiopsyllidae, which is prevalent
in fine Australo-Malayen Subregicn. An undescr-ibed spades of finis
genus, finat has been found in Middle America (Wenzel and Tipton,
_lo_g;_c;_i_t_.), is interpreted as an invader from northern South America.

2% Egtrans is another authentic South Ansrican flea which
probably first became parasitic on rodents. During the interim,
however, finis flea became more selective for larger end more adequate
hosts, sndnasism,_m_2_s_aggn_smdofinerlargemnals.

The gems M has an interesting discontinuons distribution . mile
five species are typically Neotopical end have originated in fine
Brazilian Subregicn, ane spades occurs in Baja California, in fine
Rocky Mountain Subregicn of the Nearctic Region. Bofin mining
known spades of Lung are confined to fine Manchurien Subregicn

of the Palearcu'c Region; me is present in China end fine ofiner in
China end Jenan.

It is possible finat helmet fleas of fine family Stephanocircidae
were originally exclusive parasites of marsupials (I-bpldns, 1957;
Johnson, 1957; Jordan, 1931). In America, however, they have been
secondarily adapted to rodents of the family Cricetidae, which

are more suitable hosts than marsupials. The genera Cleopsylla,

9n

Plocopsylla end Sphinctopsylla, the stephanociroid groups in Colombia,

 

are more characteristic of complex-penis type cricetine rodents. They
are prechninantly Andean fleas; however, one spades, Plocopsylla
scotinani Tipten and Mendez is known from Paneni. This fact seens to
indicate finat fine genns reached Central America in more recent times.

Didelphis mausupialis , Metachirns nudicaudatus and other
marsupials are normal hosts for Adoratopsylla intermedia c_opha,
concluded to be of South American origin. Alfinough this flea is
sanetines found on M end additional cricetine rodents, it is
evident that it has marked spedfidty for opossuns end became
associated with then throughout their evolution . Neotyphloceras
rosenbergi, another tmpical South Anerican flea, seems to be
secondarily associated with opossuns, since it has preference for a
variety of cricatine rodents, especially members of fine complex-penis
types, Oryzonys, Signodcn, Akodonn, and Thomasarys.

Rhopalopsyllidae is the most characteristic family of Colombia,
end in fine American tropics and subtropics. Evidence of host-parasite
relationships end distributions of this group suggests that it
originated in South America in association with cavicmcrph rodents.
(The latter are older. on this Continent and to which some of the
rhopalopsyllids are presently affiliated): The fossil records indicate
eifiner a South Americen origin for the cavianorphs (Hooper, 19%), or
at least invasion by their precursor-s frcnn elsewhere in the Early
Oligocene, as suggested by Patterson and Pascual (1972). If
rhopalopsyllids began as parasites of cavionorphs , later they gradually
lost their preference for these rodents and becene established on a

95

variety of cricatirne hosts , particularly of the canplex—penis-type .
The basis is finat this group of fleas does not denonstrate
significant host especificity. The large genus Polygenis , by far the
cblminant rhopalopsyllid, has experienced considerable radiation and
today more than fifty form are kncm. It appeared in some area of
fine Brazilian Subregicn, which night have well been in the territory
presently occupied by Brazil. According to Wenzel and Tipton (_l_.o_g.
£11.), endemic spades of Rhopalopsyllus and Polygenis in Middle
America may be explained on the basis of an early, pre Pleistocene,
dispersal of cricetine rodents from South America.

The most distinct exposure of endemedty in southwestern
Colanbien fleas is given by the monotypic rhopalopsyllid gents
Sgiopsyllns; it appears to be confined to certain mountains
surrounding fine Cauca Valley. Present evidence is suggestive of a
recent genesis of this flea under isolated ecological mechanisms
of mflcnown nature . The fact that Coloubian nnountains have been
insuffidently explored for manuals and fineir ectoparasites offers
the possibility finat other interesting endenic taxa remain to
be fonrnd.

96

xogm hedgehog

59m BHNNQIWQE

 

 

mesoo mod. 3%
Beam w.m_.o monogram
newsflash“ mi. @0003
moon—=06 3H? stream
Bash 5686230

 

 

 

 

 

 

 

 

 

flmoloo massage A. we «Haas 8302
as o6mfi$fi m. be Heme 828a.
goes one»? ca

mmmwnfimplu $082382

as :62m s5 Waco Hummmsnosccae
8%
anodes. £62535 m. we flame 888<

 

 

 

mam

 

. 33mg

sorghum e3 omoyge sumoflwmmmao 05 mzodnom 8:8va Gan. ems—E: 05. 6&5ng on»
5 Havana 396.30 50526.38 Eonm wagon,» can 839503 EEO so romeo we won.“ mg

a

 

mcwovHOm mam
30.386503? haw—BM

 

 

mHfiaH 0H. 602
cSaSHHooz wean»
gag 98.5

 

 

25% 8P8H§U
occHonofimo Hana

 

.5me songs
Synagogues Hangman:

mflflwgmnmfi mEmHmowa

 

 

 

 

$62388 Hesse
massage: 935

mg

 

«scare s65 can fies. Esme mo pmH A Home

97

 

 

 

 

 

 

 

 

 

 

 

.em A .28 flaws

snowshoes Hesse

minor 3
«noggin hams

 

 

wwmcwpmmfl mgwow
cooaaeom season
oHsucom sons

gunmen 96m
smog Hesse

 

38.39% moaov<
0638 haste
cope—aim Mango

 

.cm magma
ache: mommOHQe meoaox

c6683: Hesse

 

 

 

 

96958 ”60885
oneness—cosmos Hesse

ESL—22

 

"3.9.89 H manna

98

 

 

 

 

 

 

 

 

 

 

 

953838 msaawmmaoow
628 328 surgeons

$3

628 628 surgeon

 

 

é

 

gage/gowns arson?
925m. amoeba.

mHaam gala

 

859.3 and

98mm. “memo A; g

apnea Age mg

”.232:

 

"6.208 H Home

99

 

 

mmhnsmsa mzflhmmoamdoé

mgmwm mauwomo msaazmmoamdonm
959.... mHHvamsm msaHMmamHmaoé
mmhnzwsa gflwmmoaaoé

gmMm mawdmo mDHHMmAOdmmoé
959p mflnmhwsm 9323.392

 

 

 

mwmoono QHQWQQBx
mgmm gammouamq
wwomao Sahmmgwx

 

 

 

nofi £1698on

gap magmaoficmmm
gamma: mHHmwmomHo

x8 mEOwowvwfiyo
“magma memOOESomz

 

Moog wmdmwbnom

HEB EBB mummflom
£0035: Sammmomao

2&8 mflgvfi m Nady
a mgmooazmowz

93.2mm wasm
.drgw “”3358on
mmuopgvo magnum mmummnoomam

 

 

 

 

Egg madam "Sigma
383:9: Snags

 

gm

 

mkubgm gamma

 

mama flbomfl
mmvflooufimma h Emu

 

9“.va 9....va
90mg «Bhwm

meHSz haw—Emu

.mm ag

 

389% @5355.

mg

 

"8388 H mafia

100

 

 

mun—”mm mmvwamcawmoogvo

 

3mg $25002
9595 wwwnmfitwg...v gagmaoaaoé

 

mflnwm mmcflnmnaguo
madam.“ mmvflmmnfioocmuo
muflmm mmvflflmfiwmoofifio

 

M83 marsmmmaom

 

“8mg mwcmwmflom

mg

 

£58 Emma mfimm
«memo» SEE

 

Eng 5392
Panama am
03.33% 365m

“.505 868.80
Bopcwmnmowumcwo 559.5
mggm mwcwo
«mug 32mm

 

 

fig .320
9.5g Ema

ggmgm 3g
«8% figm

mg

 

 

HAULGOOv H anmB

 

XXXXXX

xxx
xx

101
X XXXX

><><><><><><><
XX XX XXNXXXX

XX

XXXXXXX
XXX ><

><><><>< XXXXXXX

 

 

 

 

 

wepuv pue
Ktuo mapuv

 

sugseq meanv
pus coon-.510
pup mg

90m tanmo

mm 01:de

mnodansoo

eopmv LEI-ROS

 

 

 

 

:33 Bowman

any—mag

H8395

Immanuel-om

 

 

hflgOH Ogv

whmummconmwm

 

L

 

 

.33“ .5938 $28 mo .55.» m5 mo 3.33%? a; .N mafia

102

 

 

 

 

3}
5%
kg 0

XXX
xxx
xx

 

 

XXXXXXXXX
.9:
- 3
§
33
8

 

 

W

[3119 ma

 

 

 

KIUO maepuv
sutseq mzemv
mmv TED-“30

20pm mnos
mm mm

 

 

WOW-0M
mnodcmsoo
4, 1’
n. .-n

8%. 3:33.—
Hmoflphboz .303ng

 

 

 

 

. $3 53.38 avg mo 38% «5 mo fiflgmfi 0% .m 33.

 

 

xxx x

103

x x xxx

xxxxxxxxx

xxx

xxxxxxxxx

xx

xxxx
xxxxxx
xxx

Qfiflfiwmmw
figmgfiagfi@fi
mgaéfimgfl

333§MHE
mfimmmwwu. E8958
ma 5mg 5x
IQ: H 933%
gagogmfieggé
fiEHBMQQQEHEfi
35383035@83£

.%.cJ§§JW., Wfi
ammwmmwflw
9 a .mwfimmm %

 

 

 

 

 

 

 

wepw pm?

At!» mapuv

“WHNWWN

 

sqnqmmw
mmanmm

 

wwwvwmmo
wammm

Immmogpq
mnmmmm

“WWW¢W8

 

 

 

mega
33ng

 

853.3

E

 

 

Immmmwmpmmn

:83 oflwuommv

mnmummaognwm

 

"GLSQ mafia

 

KEY TO THE FLE‘AS OF SOLYI'EMS'I'ERN COLQIVIBIA3

1. Thorax very reduced or compressed; antesensilial bristles
absent; females bmm into skin 2
Thorax not reduced or compressed; antesensilial bristles present;
females do not burrow into skin 3
2. Head frons angular; anterior lower margin of hind coxa with tooth-
1ike projection; not parasitic on bats.....'I\1nga penetrans
Head frons rounded; hind coxa without tooth-like projection;
parasitic on batsRhynchopsyllus pulex
3. Sword-file ridge of mesocoxa absent; mesonotum always lacking
pseudosetae; metanotum rectangular, not markedly broader
dorsally than ventrally; never more than one row of bristles
on abdmfiml terga II-VI; patch of spinelets on inside of
Sword-like ridge of mesocoxa usually present; mesonotum with or
without pseudosetae; metanotum much broader dorsdly than
ventrally; without patch of spinelets on inside of

“BEMOOOOOCOOOOOOOOOOOOOOO0.0.0.0...OOOOOOOOOOOOOOOOOOO 7

 

3

The following papers were helpful in the preparation of this key:
Johnson (195”. and 1957), Ewing (1929), and Tipton and Méndez
(1966). Rhopa10psyllus lugubris, I_2_. cacicus saevus, Polygenis
dunni, and E. roberti beebei have not been reported from
Wm Colonbia fliowever, they are likely to occur there
and are included in this key.

101$

‘4.

5.

6.

7.

8.

9.

105

Genal and pronotal comb present. . . . . . . . . . . . .Ctenocephalides £2133
Genal and pronotal comb absent...................................5
Mesothorax pleural rod present; eight tergum of female complete
dorsally; bulga of spermatheca not globular, heavily
pigmented.................................XenOpsylla chegpis
Mesothorax pleural rod absent; eight tergum of female divided
dorsally; bulga of spermatheca globular, lightly
pigmented...................................................6
Dorsal aedeagal sclerite broad throughout; crochets small and
elongate , rodlike; seventh sternum of female usually with
7-9 bristles.................................P._u_l_g)_g simulans
Dorsal aedeagal sclerite relatively long and slender; crochets
expanded apically, not rodlike; seventh sternum of female
usually with "-5 bristles.....................§_L_11_e3_< irritans
Head possessing true helmet; ctenidia aheays present on helmet,
gena and pronotum.........-..................................8
Head without helmet; ctenidia absent or present, limited to gena
and/or pronotumu..........................................12
Helmet separated from head dorsally; vertical comb
posteromarginal.............................................9
Helmet not separated from rest of head; vertical comb mesad....
........................................Clqusylla monticola
'IWo large genal bristles anterior to cibarial pump; eighth
tergum of male with apodeme; female spermatheca barrel-shaped,

without internal tubercle , and hilla not enlarged

basally.00......OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO.0.00.00.00.10

10.

12.

13.

106

One or two large genal bristles posterior to or vertically in line
with cibarial pump; eight tergun of male lacking apodeme;
female spermatheca not barrel-shaped, with internal
tubercle orwith hilla enlargedbasally...................11

Genal canb teeth reduced, slightly longer than
broad..Plocopsylla phyllisae

Genalcmbteethnomel,mbrethanmiceas long
as broadPlocopsyllathor

Pronotal comb spines nmbering about 18, not pointed; male with
vertical row of four close-set spiniforms on seventh
sternum; female with subcaudal row of heavy spiniform
setaecnninth stemmwmwmunctopsylla diomedes

Pronotal comb spines nmbering about 30, pointed; male only with
normal bristles on seventh sternum; female without subcaudal
row of heavy spiniform setae on ninth
sternunSphinctopsylla tolmere

Anterior margin of head conical, having two short spiniform
bristles; tibiae with dorsal curb of short, stout
bristles"...............................Leptopsy11a segnis

Anterior margin of head not conical, without short spiniform
bristles; tibiae without chrsal curb of short, stout
bristles..................................................l3

Severelabdafinaltergawith well developed combs..............ll4

Abdmfimltergawithout curbs”................................15

1"".

15.

16.

17.

18.

1E3.

107

Crochets with convex margin oriented dorsad; female seventh
sternum with caudal margin tnmcate.......Ctenidiosomms rex
Crochets with convex margin oriented ventred; female seventh
sternumwith caudalmargnnottnmcate
Ctenidiosomus treubi
Anteroventrel margin head with two large genal spines; found on
batsSternopsylla distincta speciosa
Anteroventrel margin of head with more than two or lacking genal .
spines; not foundcnbats.................................16
Having a combination of the following characters: anterior
tentorial arm present , always inserted anterior to eyes;
mesopleurel rod not dorsally bifuroated; ventral margin of
pronotum not bilcbed; fifth tarsal segment always with four
pairs of plantar bristles.................................l7
Laddngabove ccmbinatim of charectersBl
Antennal club symmetrical; without apical spinelets
onmetanotum.............................Tetrepsyllus comis
Antennal club asymmetrical; apical spinelets present on
metanotum.................................................18
Prontoclypeal margin of head subconical; pronotal curb
present.............................Scolopsy]_lus colombianus
Prontoclypeal margin of head romded; pronotal comb

ammtOOO...OOOIOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOCOOO00.019

Prosternosame projecting downward between coxae; mesocoxa

0.00.0.0...OOOOOW®YIIIJSCOOOOOOOO0.0.0....000000000020

108

Prosternoscme not projecting downward between coxae; mesocoxa
asymmetrical, obviously broadest basally
PolygenisuZZ

Spiracle of metepimere oblong, prolonged dorsally; bulga of
spermatheca globular.................Rhopalg>syllus lugubris

 

mt glObWOOOOOI.0..0....0..OOOOOIOOOOOOOOOOOOOOIOO0.0.0.21

Labial palpus reaching apex of coxa I or beyond; movable process

shaped......................Rhopalopsyllus cacicus saevus

 

Labial palpus not reaching apex of coxa I; movable process of
clasper about as long as distal arm of ninth sternum;

smtbem smy S- ShameOOOO0.000000000000000000000000
0.0.0.0000...OOOOOOOOOOOOOCOmom-opsynm ammis tllpinm

 

Distal arm of ninth sternum much shorter than proadmal arm; heel
of ninth sternum strongly angular; inner tube of aedeagus
reflexed dorsally, not coiled apically.....................23

Distal arm of ninth sternum about equal length or longer than
proadmal arm; heel of ninth sternum weakly angular; inner

apiaIlYIOOOOOOOOOOOOOOOIOOOOOOOOIOOOOOCOOOOIOOOOOOOOOOOOO0.2.4

 

20.
Spirecle of metepimere rounded or ovoid; bulga

21.
of clasper longer than distal arm of ninth sternum;
spermatheca somewhat boomerang-

22.
tube of aedeagus reflexed ventrally, coiled

I;

Females of Polygenis are difficult to identify and the characters

used in this key pertain almost entirely to males. The females
of E. tra idoi, n. sp. and g. hom‘ i n. sp., are unknown.

23.

21$.

25.

26.

27.

109

Clypeal tubercle above eye level; posterior margin of fixed
process convex; spermatheca sinuate , without separation
between bulga and hilla..................Polygenis thurmani

Clypeal tubercle at eye level; posterior margin of fixed process
sinuate; spermatheca humped, with distinct separation
between bulga and hilla. . . . . . . . .Polygenis klagesi samuelis

Distolateral lobes of aedeagus with angular projection; eight
sternum of male divided in half ventrally, with short

dorsocaudal extensim.............Polygenis trapidoi, n. Sp.

 

Distolateral lobes of aedeagus without angular projection;
eight sternum of male not divided in half ventrally,
without dorsocaudal extensronZS
Apex of distal arm of male ninth sternum bearing distinct group
of stout bristles....................Polygenis bohlsi bohlsi
Apex of distal arm of male ninth sternum not bearing distinct
group of stout bmstlesZB
Aedeagal fender present; dorsal margin of fixed process of clasper
sinuate; spermatheca with cribose bulga, its ventral margin
interrupted at bulga-hilla junction (with probable exception
of 1:. Mi, n. sp)27
Aedeagal fender absent; dorsal margin of fixed process of clasper
usually slightly convex; spermatheca not cribcse, its ventral
margin continuota at bulga-hilla junction
Polygenis pradoi
Aedeagal side piece above basal part of inner tube; aedeagal ribs

not nmerous......................Polygenis delpontei n. sp.

28.

29.

30.

31.

32.

110

Aedeagal side piece absent or below basal part of inner tube;
aedeagal ribs very numerous28

Aedeagal lateral lobes very reticulate; subapical ridge of median
dorsal lobes of aedeagus present

...............................Polygenis caucensis n. sp.

 

Aedeagal lateral lobes not reticulate or faintly reticulate;
subapical ridge of median dorsal lobes of aedeagus
absent.....................................................29

labial palpus extending to trochanter I; several distal arm
bristles of ninth sternum very long, approximately four
times maxiJmmm width of distal arm

0.0.0.0....0.......0....0.00000000000000000PoygmiS dmmj-

 

labial palpus not extending to nochanter I; distal arm bristles

of ninth sternum short or moderate length , never very

10ng.......................................................30
Aedeagal fender well developed, half-moon shaped; distal arm

of ninth sternum distinctlybroadmedially..................

Polygenis roberti beebei
Aedeagal fender reduced to slender, arched, inconspicuous

structure; distal arm of ninth sternum not ch'stinctly

broad medially.................Polygenis hgpkinsi, n. sp.
Genal cambpresent....32
Genalcombabsent...............................................33
First spine of genal comb almost overlapped by second; trabecula

centralis present; labial palprs S-segmented; both sexes

111

with three antesensilial bristles............................

0.0.0.0000...OOOOOOOOOOOOOOOOOOOOONeOtyphJ-mems mam

 

First spine of genal comb rot overlapped by second; trebecula
centralis absent; labial palpus lt—segmented; both sexes with
mmteserBj-lial bfi-StleSOOQCOCCIOOOOOOOCOOCOOOOOOOOOOOOOOOO

.Adoratopsylla intermedia copha

 

33. Pronotal comb having more than twenty four spines; bristles of
second antennal segment long in both sexes...................

.Dasypsyllus gallinulae perpinnatus

 

Pronotal comb usually with less than twenty four spines; bristles
of second antennal segment short in male, never reaching
apex of club in female3|+

3t}. Ptotibia with seven dorsal notches containing paired bristles;
meso- and metatibia with six dorsal notches containing
paired bristles proximal to only single dorsal
bristle...”...............................Pleochaetis smiti

Protibia with five or six dorsal notches containing paired
bristles; meso- and metatibia with five dorsal notches
containing paired bristles proximal to only single dorsal

bfistle...0.000COOOOOOOOOOOOOOleetiS ”La-toms. amms

Figure 6. General @pearance of female of Ctenocephalides

felis (Bouche) .

List of Abbreviations

Abbreviation Explanatim

AN.

we?
OSDU)
a

["1 00
CUE-.0
%

. mm.
9? .

Farrrm
wram
s

maso

U)
o

museum
%

Antenna

Antesensi lial seta
Anal stylet of female
Bursa copulatrix
Coxa

Claw of tarsus
Dorsal anal lobe
Bye

Epipharynx

Femur

Prons

Genal ctenidium
Lacinia

Labial palp
Lateral plantar bristles
Maxillary lobe
Maxillary palp
Mesepimere
Mesepisternum
Mesonotum
Metepimere
Metanotum
Metasternum
Occiput

Ocular bristle
Pronotum

Prmotal ctenidium
Proepimere
Sensilium
Spermatheca
Sternum

Tergum

Tarsus

Tibia

Ventral anal lobe

112

:_> Hm: I I

=>.P

.).h

Iago-C

.m gum

    

 

 

Figure 7. Structures of apex of aedeagus of Scolopsyllus

colombianus Mendez .

 

List of Abbreviations

 

 

Abbreviation Explanatim
A.M.S. Apico-median sclerite
AP.S. Apodemal strut
CR. Crochet
Cr.P. Crochet processes
C.S. Crescent sclerite
DL. L. Distolateral lobes
F.M. Fluted membrane
H.L. Heel at base of aedeagal poud'r
I.T.-A Apical portion of sclerotized
inner tube
I.T.-B Basal portion of inner tube
L.L. Lateral lobes
M.D.L. Median dorsal lobes
P.R. Penis rods
PS.T. Pseudotube

113

DL. L.

 

Figure 7.

Figure 8

Neotyphloceras rosenbergi (Rothschild)

Female

A. Head, prothorax and procoxa.
B. Modified abdominal segments.

C . Spermatheca.

 

114

Figure 8.

Figure 9

Adoratgpsylla intermedia flha Jordan

Male

Head, prothcrax and procoxa.
Cflnitaliao
Apex of aedeagus .

Female

Spermatheca and 7th abdominal segment.

115

 

 

5. 4.4m

Figure 9.

COED>

Figure 10

Ctenidiosomus traubi Johnson

Male

Head, prothcrax and procoxa
Ninth sternum.
Clasmro

Apex of aedeagrs .

116

 

Figure 10 .

Figure 11

Cleopsylla monticola Smit

Male

A. Head, prothcrax and procoxa.

B. Mesothcrax, metathorax and tergum I.

117

 

Figure 11 .

Figure 12

Sphinctopsylla diomedes Johnson

Female

A. Head, prothcrax and proccm.
B. Modified abduminal segments.
C. Spermatheca.

118

 

Figure 12 .

005:!)

Figure 13

PlocOpsylla phyllisae Smit

 

Male

Head, prothcrax and fore coxa .
Mesothorax, metathcrax and tergum I.
Modified abdomu'nal segments .
Spermatheca.

119

 

Figure 13.

Figure l”.

Plocopsylla thor Johnscn

Male

A. Head, prothcrax and fore coxa.

120

 

 

 

 

 

 

 

unpu>

Figure 15

Sternopsylla distincta speciosa Johnson

 

Male

I-had, prothcrax and procoxa.
Eight sternum.
Process and movable finger of clasper.

Distal arm of ninth sternum.

Female

Modified abdominal segrrents .
Aral stylet and ventral anal lobe.
Spermatheca.

121

 

 

 

Figure 15 .

coyote

Figure 16

Leptopsylla segpis (Schonherr)

Male

Head, prothcrax and fore coxa.
Mesothorex, metathorax and tergum I .
Process and movable finger of clasper.

Hind femur and hind tibia.

Female

Modified abdominal segments .
Spermatheca.

122

 

 

 

Figure 15.

Figure 17

Dasypsyllus gallinulae perpinnatus (Baker)

Male

A. Head, prothcrax and procoxa.
B. Genitalia.

Female

C . Spermatheca and 7th abdxminal segmert.

123

 

Figure 18

Pleochaetis smiti J ohnscn

 

Male

Head, prothcrax and procoxa.
Process and movable finger of clasper.

Distal arm of 9th sternum.
Female

Modified abdominal segments .
Spermatheca.

121$

 

Figure 18 .

Figure 19

Tetrapsyllus comis Jordan

Female

A. Head, prothcrax and procoxa.
B. Modified abdominal segments.

C. Spermatheca.

125

 

 

Figure 19 .

Figure 20

Polygeris bohlsi bohlsi. (Wagner)

Male

Head, prothcrax and fore coxa.
Process and movable finger of clasper.

Apex of aedeagus.
Female

Distal arm of 9th sternum.

Spermatheca.

126

 

Figure 20 .

Figure 21

Polygeris caucensis, n. sp.

Male

A. Head and prothcrax.
B. Process and movable finger of clasper.

C. Modified abdominal segments.

127

 

 

at

Figure 21.

Figure 22

Polygenis cauceisis, n. sp.

Male

A. Distal arm of 9th sternum.

B. Apex of aedeagus.

Ferale

C. Spermatheca.

128

 

Figure 22 .

Figme 23

Polygenis delpontei, n. sp.

Male

A. Head and prothcrax.
B. Process and movable finger of clasper.
C. Modified abdominal segments.

129

 

 

.--—_
_—-‘L‘
a‘,‘
—

          

Figure 23.

Figure 2t}

Polygenis delpontei, n. sp.
Dale

A. Distal arm of 9th sternum. .

B. Apex of aedeagra.

130

 

.‘r
””133". . , '
"‘v _"'34;51-

I
O
' I
I
' l
‘ I
l
1..
r
l
.93.!
.....,.
'. I"?
‘1 r". ‘
I
o
i
O
53"
. u»...
a"
‘ A
8
u
U
A;

   
  
         
     

  

 
 
         
 

'.o.'-.o ‘ 00 o:-'-:.‘::::':' '
o c
a.
o
- p
o 0' ..._-_: -
" ,_.;::a;fii':§‘-"
. .fiau
( no". 04-.Q ‘3‘ = ‘
r s

Figure 2a.

00513)

Figure 25

Polygeris delpontei , n . sp .

Ferale

Head and prothcrax.
Modified abdominal segme'rts.
Dorsal and anal lobes of proctiger.

Spermatheca.

131

 

 

 

Figure 25.

Figure 26

Polygenis dunni (Jordan 8 Rothschild)

 

Male

A. Head, prothcrax and procoxa.
B. Genitalia.

Female

C. Spermatheca and '7th abdaminal segment.

132

 

Figure 26 .

Figure 27

Polygeiis hgphnsi, n. sp.

 

Male

A. Head and prothcrax.
B. Process and movable finger of clasper.
C. Modified abdominal segments.

 

 

 

 

 

 

Figure 28

Polygenis hopkinsi , n. sp.

Male

A. Distal mm of 9th sternum.

B. Apex of aedeagus.

   

0

 

-u.. ..u'
...-....

  

131+

 

he ....
. .
. m ..
.. ..t
. .
... a ...
o I.

.I
3‘30- 4?

 

Figure 29

Polygenis klagesi (Rothsd'rild)

 

Male

A. Head, prothcrax and procoxa.
B. Genitalia

Female

C. Spermatheca and 7th abdominal segment.

135

 

 

 

Figure 29 .

Figure 30

Polygenis pradoi (Wager)
Male

Head, prothcrax and fore coxa.
Process and movable finger of clasper.

Apex of aedeagus.
Distal arm of 9th sternum.

Female

Spermatheca.

136

 

Figure 30 .

Figure 31

Polygenis roberti beebei (I. Fox)

 

Male

A. Head, prothcrax and procoxa.
B. Geiitalia.

Ferale

C. Spermaflreca and 7th Abdominal segment.

137

 

Figure 31 .

00511?»

Figure 32

Polygenis thurmani Johnson
Male

Head, prothcrax and fore coxa.
Process and movable finger of clasper.

Apex of aedeagus.
Distal arm of 9th sternum.

Female

Spermatheca.

138

 

......vnilff

‘7'? so

 

 

U
u: t .
.. $3.32», »

,9"

r t '.-'

../.
' ..
.
. .
J D I
u
.
O Q. .
4 $3....
0 q.
4 o.
n .
. aw.
J».
. :.
....1
,\
... .I
o
. t
.
.1
.fi
h
a
Q. 5
.. s

 

Figure 32 .

Figure 33

Polyng trapidoi, n. sp.

 

Male

A. Head, prothcrax and procoxa.
B. Process and movable finger of clasper.

C. Modified abdominal segments.

139

 

    
    
 

. .....
§
'9‘
“a
' . --
"3——-.r:.u.-.: - '2.
.N.\ """-~r.r& up
.“
2,.
-\.

I
a

in
" ,’ .r
O o ____
. <‘
x |
' —— 1“"
t , Js’
' C

Figure 33.

Figure 3a

Polyggr__ris idoi , n. sp.
Male

A. Distal arm of 9th sternum.
B. Apex of aedeagus.

1'40

 
   

 

Figure 31;.

COED>

Figure 35

Scolopsyllus colarbianus Mendez

Male

Head, prothcrax and procoxa.
Mesothorax, metathorax and tergum I .
Process and movable finger of clasper.

Distal arm of 9th sternum.

1'41

 

 

 

 

 

Figure 35 .

Figure 36

Scolopsyllus colcmbianus Merdez

Female

Head and prothcrax.
Modified abdominal segments.
Dorsal and anal lobes of proctiger.

Spermatheca.

 

 

 

 

 

 

 

Figure 36.

Figure 37

RhgpalOpsyllus austrelis tupinus Jordan 8 Rothschild

 

Male
A. Head, prothcrax and procoxa.
B. Genitalia.

Ferale

C. Spermatheca and 7th abdominal segment.

1&3

 

 

 

 

 

 

 

 

Figure 37.

Figure 38

Rhopalopsyllus cacicus saevus Jordan 8 Rothschild

Male

A. Head, prothcrax and procoxa.
B. Genitalia.

Ferale

C. Spermatheca and 7th abcbmuinal segment.

114M

 

 

 

 

 

 

 

 

 

Figure 38.

Figure 39

Rhopalopsyllus lugubris Jordan 8 Rothschild

Male

A. Head, prothcrax and procoxa.
B. Genitalia.
C. Process and movable finger of clasper.

Ferale

D. Spermatheca and 7th abdominal segment.

1145

 

 

Figure 39 .

Figure 140

Xenogylla cheopis (Rothschild)

Male

Head, prothcrax and procoxa.
Parocess and movable finger of clasper.

Distal am of 9th sternum.

Ferale

Modified abcbmuinal segments .
Spermatheca.

1'46

 

Figure 90 .

Figure Ml

Ctenocephalides felis (Bouche)
Male

A. Head, prothcrax and procoxa.
B. Genitalia.

Ferale

C. Spermatheca and 7th abdominal segmert.

197

 

Figure 1&2

Pulex irritans Linnaeus

Male

Head, prothcrax and procoxa.
Detached process and movable finger
of clasper.

Distal arm of 9th sternum.

Ferale

Modified abdum'nal segments .

Spermatheca.

198

 

Figure #2 .

"1157100

Figme 1+3

Rhynchopsyllus pulex Haller

Male

Head, prothcrax and procoxa.
Mescthorax, metathorax and lst abduminal

segment.
Modified abdominal segments.
Process and movable finger of clasper.
Distal arm of 9th sternum.

Apex of aedeagus .
Female

Spermatheca and modified abcbminal segments .

1149

 

Figure R3.

Figure 141+

'Iunga penetrans (Linnaeus)

 

Male

A. Head, prothcrax and procoxa.
B. Geritalia.

Female

C . Spermatheca and 7th abdominal segrent .

150

 

Figure nu .

BIBLIOGRAPIH

BIBLIOGRAPHY

Aller, J. A. 1912. Mammals frumwestern Columbia. Bull. Amer.
Mus. Nat. Hist. 31(14): 71-95.

. 1913. New mammals from Colombia and Ecuador. Amer.

. 1915. New South American Mammals. Bull. Amer. Mus.

. 1916. List of mammals collected in Columbia by the
AmEican Museum of Natural History expeditims. 1910-1915.
Bull. Amer. Mus. Nat. Hist., 35(18): 191-238.

Baker, R. H. 1967. Distribution of recent mammals along the
Pacific Coastal Lowlands of the Western Hemisphere.
Systeratic Zoology 16(1): 28—37.

Beek, K. J. and D. L. Bramao. 1968. Nature and Geography of South
American Soils. Pages 82-112 _i_n_ E. J. Fittkau, J. Illies,
H. Klinge, G. H. Schwabe and H. Sioli, eds. Bibliography
and Ecology in South America. Vol. 1. Dr. w. Junk N. V.
Publishers. The Hague.

Borrero, J. I. 1967. Mamiferos Neotropicales. Universidad del
Valle, Colurbia. 190 pp.

Biirgl, H. 1961. Historia Geolégica de Colurbia. Rev. Acad. Col.
Cienc. Bx., Fis. y Nat. 2: 137-191.

Cabrera, A. 1958, 61. Catalogo de los mamfiferos de America del Sur.
Rev. Mus. Argentino Cienc. Nat. "Bernardiro Rivadavia."
IV(1) 1958: i-iv 3‘ 1-308, (2) 1961: v-mdi ’l 309-732.

, and J. Yepes. 19am. Mamiferos Sudamericanos (Vida,
sturbres y Descripcién). Historia Natural Bdiar,
Compafiia Argentina de Editores, Buenos Aires. 370 pp.

Cali. Virus Laboratory. 1969 Report. Cali, Columbia:
Universidad del Valle, 1965. 52 pp.

Cali Virus Laboratory. 1967 Report. Cali, Columbia: Universidad
del Valle, 1968. 155 pp.

151

152

Cali Virus Laboratory. Cumbined 1968, 1969 Reports, Cali, Columbia:
Universidad del Valle, 1970. 209 pp.

Chabaud, A. G. 1997. Les arthropodes vecteurs de la peste
bubonique. Ann. de Parasitologie 22(3-9): 192-200.

Chapman, F. M. 1917. The distribution of bird-life in Columbia:
A contribution to a biological survey of South America. Bull.
Amer. Mus. Nat. Hist., 36. 729 pp.

Clay, T. 1951. The Mallophaga as an aid to the classification of
birds, with special refererce to the structure of feathers.
Proc. Xth Internatl. Ornith. Cong., pp. 207-215.

Costa Lima, A. da and C. R. Hathaaay. 19'46. Pulgas: Bibliografia,
catalogo e animais por elas sugados. Mumgafias do
Instituto Oswaldo Cruz u. 522 pp.

Dunn, L. H. 1929. Notes on some insects and other arthropods
affecting man and animals in Colombia. Amer. Jour. Trop.
Med. 9: “93-508.

Eidt, R. C. 1969. The climatology of South America. Pages 5u—81
inE.J. Fittkau,J.I11ies, H. Klinge, G. H. SchwabeandH.
8101i , eds . Biogeography and ecology in South America,

Dr. W. Jurk N. V. Publishers. The Hague.

Fspinal T., L. S. 1968. Vision Ecolégica del Departamento del
Valle del Cauca. Universidad del Valle. 105 pp.

, y B. Murtenegro. 1963. Formaciones vegetales de
Columbia. Memoria explicativa sobre el mapa ecolégico. Imp.
Canal Ramirez, Bogota. 201 pp.

Being, H. B. 1929. A Manual of External Parasites. Charles
C. Thumas, Publisher, Baltimore, Md. 225 pp.

FAQ-UNESCO. 1971. Soil map of the World. Volure IV. South America.
Unesco. Paris.

Fittkau, E. J. 1969. -The faura of South America. Pages 6214—658
in E. J. Fittkau, J. Illies, H.1<1inge, G. H. Scheme and H.
Sioli, eds. Biogeography and ecology in South America. Vol. 2 .
W. Junk N. V. Publishers. The Hague.

Fuller, H. S. 191:2. Notes on Neotropical Siphonaptera. Revista de
Enturologia 12: 39-91:.

Gast Galvis, A. 1950. Lista de la pulgas encontradas en Columbia.
Revista de Hig'ene 29: 195-199.

153

Gyldelstolpe, N. 1932. A Mantel of Neotropical Sigmodon Rodents.
Kungl. Svenska Veterskapsakademiens . Handlingar (3)XI,
Mo 3: 1.16.4.

Haffer, J. 1967. Speciation in Colombian forest birch west of the
Andes. Amer. Mus. NomNurber 2291:: 1-57.

Harm-en, T. V. D. 1961. The Quarternary climatic changes of northern
South Arrerica. Ann. New York Acad. Sci., 95: 676-683.

Hershkovitz, P. 19'41. The South American harvest mice of the genus
Reittuodurturrys. Occ. Pap. Mus. Zool. Univ. Mich mm: 1-7.

. 1997. Mammals of Northern Columbia, Heliminary report
No. l: Squirrels (Sciuridae). Proc. U. 8. Nat. Mre.,

. 1998. Mammals of Northern Colombia, Preliminary report
No. 2: Spiny rats (Echimyidae), with supplemntal notes on
related forms. Proc. U. 3. Nat. Mus., 97: 125-190.

. 191:9. Mammals of Northern Colombia, Preliminary
report No. 5: Bats (Chiroptera). Proc. U. 8. Nat. Mus.,
99: 1429-4451:.

. 1958. A geographical classification of Neotropical
mammals. Fieldiana: Zool., 36: 581-620.

. 1960. Mammals of Northern Colombia, Preliminary
report No. 8: Arboreal rice rats, a systeratic revisim
of the subge'rus M, gems M. Proc. U. S. Nat.
Mus., 110: 513-568.

. 1962 . Evolution of Neotropical cricetine rodents
(Mt—Jridae) with special reference to the phyllotine group.
Fieldiana: Zool., #6. 52% pp.

. 1966. Mice, land bridges and Latin America faunal
1nterchange. Pages 725-7147 in_ R. L. Welzel and V. J. Tipton,
eds. Ectoparasitcs of Panama. Field Mus. Nat. Hist.,
Chicago, Illinois.

 

. 1969. The evolution of mammals on southern continelts.
. The recent mammals of the Neotropical Region A
zoogeographic and ecological review. Quart. Rev. Biol. ,
”'4: 1.700

151+

. 1972. The recent mammals of the Neotropical Region: a
zoogeographic and ecological review. Pages 311-1431 _ifl A.
Keast, F. C. Erk and B. Glass, eds. Evolution, Mammals and
Southern Continerts . State University of New York Press ,

Albany, N. Y.

Irbldridge, L. R. 190.7. Determination of world plant formations from
simple climatic data. Science 105: 367-368.

Holland, G. P. 1961:. Evolution, classification and host—relationships
of Siphonaptera. Ann. Rev. Ent. 9:123-1146.

Hooper, E. T. 19lt9. Faunal relationships of recent North American
rodents. Misc. Pub. Mus. Zool. Univ. Mich. 72: 1—28.

Hopkins, G. H. B. 19142. The Mallophaga as an aid to the classification
of birds. Ibis 814: 914-106.

.. 1957. Host-associations of Siphoraptera. Pages Git-87 in
net symposium of host specificity among parasites of
vertebrates . Neuchatel .

, and M. Rothschild. 1953. An illustrated catalogue of the
Rothschild collection of fleas (Siphonaptera) in the British
Museum (Nat. Hist.). Vol. I. Turgi.dae and Pulicidae. British
Museum (Nat. Hist.), xv # 361 pp.

, and M. Rothschild. 1956. An illustrated catalogue of the
Rothschild collectiur of fleas (Siphoraptera) in the British
Museum (Nat. Hist.). Vol. II. Copt0psyllidae, Vermipsyllidae,
Steptanociroidae , Is d'rnopsyllidae , Hypsophthalmidae , and
Xiphiopsyllidae. British Mtseum (Nat. Hist.), xi I 1:145 pp.

, and M. Rothschild. 1962. An illustrated catalogue of the
Rothschild collection of fleas (Siphonaptera) in the British
Museum (Nat. Hist.). Vol. III. HystriChopsyllidae
(Acedestiinae, Aromiopsyllinae, Hystrictopsyllinae ,
Neopsyllinae , Rhadinopsyllirae and Steopcniinae) . British
Museum (Nat. Hist.), viii ’4 560 pp.

, and M. Rothschild. 1966. An illustrated catalogue of the
Rothschild collection of fleas (Siphoraptera) in the British
Museum (Nat. Hist.). Vol IV. Hystrichopsyllidae
(Ctenophthalmrinae , Dinopsyllinae , Doratopsyllinae and
Listropsyllinae). British Museum (Nat. Hist.), viii # 5149 pp.

, and M. Rothschild. 1971. An illustrated catalogue of.the
Rothschild collection of fleas (Siphonaptera) in the British
Museum (Nat. Hist.). Vol. V. Leprtopsyllidae and
Ancistropsyllidae. British Museum (Nat. Hist. ), viii # 530 pp.

155

Jacobs, 0., H. Biirgl and D. L. Conley. 1963. Backbone of Colurbia.
Am. ASSOC. P612101. @601. M81110, 2: 62-720

Johnson, P. T. 19514. Notes on Pleochaetis Jordan, 1933, from
Columbia, with the description of a new species (Siphonaptera:
Ceratcphyllidae). Jour. Wash. Acad. Sci., um: 289-296.

 

. 1957. A Classification of the Siphonaptera of South America.
Mano mt. SOC. waSho 5: 1-299, 11‘“ p180

Keast, A. 1968. Evolution of mammals on southern continents as
backgrounds for mammalian evolution. Quart. Rev. Biol. 13(3);
225-233.

Ku'rten, B. 1969. Continental drift and evolution. Sci. Amer.
22: su-su.

Lewis, R. E. 1972. Notes on the geographical distribution and host
preferences in the order Siphonaptera. Part 1. Pulicidae.
J. Med. Brit. 9: 511-520.

. 1973. Notes on the geographical distributiu'r and host
preferences in the order Siphoraptera. Part 2. Rhopalopsyllidae,
Malacopsyllidae and Vemipsyllidae. J. Med. Ent. 10: 255-260.

 

. 19 7”. a. Notes on the geographical distribution and host
preferences in the order Siphonaptera. Part 3 .
Hystrichopsyllidae. J. Med. Bnt. 11: 1u7-167.

. 1971: b. Notes on the geographical distribution and host
preferences in the order Siphmaptera. Part I: . Coptopsyllidae ,
Pygiopsyllidae, Stephanociroidae and Xiphiopsyllidae. J. Med.
But. 11: ”03-013.

. 19 7a c. Notes u: the geogaphical distribution and host
preferences in the order Sipronaptera. Part 5.
Ancistropsyllidae, Chimaeropsyllidae, Isdmopsyllidae,
Leptopsyllidae and Macropsyllidae. J. Med. Bnt. 11: szs-suo.

. 1975. Notes on the geographical distribution and host
prefererces in the order Siphonaptera. Part 6. Ceratcphyllidae.
J. M. Brit. 11: 658-676.

Louris, F. G. 1911:. Origin of South American Fauas. Bull. Geol. Soc.

Macchiavello, A. 19148. Siphu'raptera de la Costa Sur-Occideital de
América (Primera lista y distribuoién zoogeografica). Bol.
Ofic. Sanit. Paramer. 27: 1:12-1469.

156

. 195m. Reservoirs and vectors of plague. J. Trop. Med.
Hyg., 57: 3-8, '#5448, 65—68, 87-9'4, 116-121, 139-196, 158-171,
191-197, 220-22'4, 238-293, 275-279, 29':-298.

Méndez, E. 1968. Scolopsyllus columbianus, new genus and species of
the family Rhopalopsylfldae (Siphoraptera) from Columbia. J .
Med. Bnt. 5(3): l405-1110.

, and H. Hansser. 1975. A new Kohlsia from the Republic
of Columbia (Siphonaptera: Ceratcphyllldae) . Proc. Ent. Soc.

Moll, A. A. and O'Leary, S. B. 191:5. Plague in the Americas. Pan
Amer. San. Bur. (Wash., D. C.) Publ. 225: 1-280.

Muller, P. 1973. The dispersal centres of terrestrial vertebrates
vertebrates in the neotropical realm. Bicgeografica . Vol . II .
W. Julk N. V. Publishers. The Hague. 22': pp.

Nygren, W. E. 1950. Bolivar geosyncline of northwestern South America.
Bull. Am. Assoc. Petrol. Geol., 3n: 1998-20006.

Olssur, A. A. 1956. Columbia. Pages 295-326 i_n_W. F. Jenks, ed.
Handbook of South American Geology. The Geological Society of
America Meroir 65, pp. 295-326.

Osgood, W. H. 1912. Mammals from western Verezuela and eastern
Columbia. Field Mus. Nat. Hist., 2001. Ser., 10(5): 33-66.

. 1993. The mammals of Chile. Field Mus. Nat. Hist.,
2001. Ser., 30: 1-268.

Patiflo-Camargo, L. 19140. Artropocbs hematéfagos de la fauna
colombiana. Rev. Acad. Colomb. Cienc. Exact” Fis. y Natur.

3: 337-38“.

Patterson B. 1957. Mammalian phylogery. Pages 1548 1lr_r_ E. Mayr and
J. Baer, eds. Premier Symposium sur la Specific1té Parasitaire
des Parasites de Vertebres . Paul Attinger, Neuchatel .

, and R. Pascual. 1968. Evolution of Mammals on Southern
Gtinelts. V. The Fossil Mammal Faua of South America.

Pollitzer, R. 195'4. Plague. hbrld Health Organization Morograph
Swies NO. 22, Pp. 1‘6980

Raven, P. H. and I. Axelrod. 1975. History of the flora and fauna
of Latin America. Amer. Sc. 63: u20—u29.

157

Reljifo-Salcecb, S. 1991+. Notas ertcmolégicas regiuales. Pages
13443 i_n_ Cespedesia 3(9-12), 197'}. Thesis.

Sanchez E., H. 1965. Elerentos de geografia geleral de Columbia.
Editorial y tipografia Bedout, Medellin, Colombia. 222 pp.

Sauer, C. O. 1950. Geography of South America. Pages 319-31.“: i_n_
J. H. Steward, ed. Handbook of South American Indians. Vol. 6.
Smiths. Inst. Bureau of Amer. Ethnol. Bull. 1'13.

Savage, J. M. 1979. The Isthmian link and the evolution of
Neotropical mammals. Contr. Sci. Nat. Hist. Mus. Los Angeles

Scott, W. B. 1937. A History of Land Mammals in the Western
Hemisphere (Revised Editiur). The Macmillan 00., N. Y. 796 pp.

Simpson, G. G. 1990. Review of the mammal-bearing tertiary of South
America. Proc. Amer. Phil. Soc. 83(5): 6'49-709.

. 19143. Turtles and the origin of the fauna of Latin
'ca. Amer. Jour. Sci. 2141(7): l:13-1129.

. 190.5. The principles of classificatiur and a classificatiul
o mammals. Bull. Amer. Mus. Nat. Hist., 85: i-xvi 4 1-350 pp.

. 1950. History of the faura of Latin America. Bull. Amer.

. 1969. South Arrerican Mammals. Pages 879-909 _1n_ E. J.
Flttkau, J. Illies, H. Klinge, G. H. deabe and H. Sloli, eds.
Vol. 2. Dr. W. Julk N. V. Publishers. The Hague.

Stirtcn, R. A. 1950. Late Cenozoic avenes of dispersal for
terrestrial animals between North America and South America.
B1111. @10 SOC. Al“. 61: 15'41-15'42.

. 19 5 3 . Vertebrate paleontology and curtinental
stratigraphy in Columbia. Bull. Geol. Soc. Am. 6'4: 603-622.

Tate, G. H. H. 1931. Random observations on habits of South American
mammals. J. Mammal. 12: 2'48-256.

. 1932 a. The taxonumic history of the South and Central
American cricetid rodents of the gems m. Part 1:
Subge'rus m. Am. Mrs. Nov. 579: 1-18.

. 1932 b. The malaria history of the South and Central
'can cricetid rodents of the genus m. Part 2:

SW Ohm, Thalluryscre, and I‘M. Am. Mus.

158

. 1935. The taxonomy of the genera of Neotropical
Hystricoid rodents. Bull. Am. Mus. Nat. Hist. 58: 295-997.

Tipton, V. J. and C. E. Machado-Allison. 1972. Fleas of Venezuela.
Pages 1-115 2:“, Ectoparasites of Venezuela. Brigham Young
University Solence Bulletin. Biol. Ser. 17.

, and E. Méndez. 1966. The fleas (Siphoraptera) of Panama.
Pages 289-386 in R. L. Wenzel and V. J. Tipton, eds.
Ectoparasites of Panama. Field Mus. Nat. Hist., Chicago,
Illinois.

Tamsitt, J. R. and I. Fox. 1970. Records of bat ectoparasites from
the Caribbean region (Siphonaptera, Acarina, Diptera). Canad.
Jour. Zool. '48: 1093-1097.

Vanzolini, P. E. 1973. Paleoclimates, relief, and species
multiplicatiorn in Equatorial forests. Pages 255-258 in B. J.
Meggers, E. s. Ayersu, and w. D. Duckworth, eds. Tm‘p‘i’cal
forest ecosystems in Africa and South America: a curparative
review. Smithsonian Institution Press , Washington.

Valentine, J. W. and E. Moores. 1979. Plate tectonics and the
history of life in the oceans. Sci. Amer. 230: 80-89.

Vuilleumier, B. S. 1971. Pleistocene changes in the faua and flora
of South America. Science 175: 771-780.

Weeks, L. G. 19'47. Paleogeography of South America. Bull. Amer.
Ass. Petrol. Geol. 31: 119'1-1291.

Wenzel, R. L. and V. J. Tipton. 1967. Some relationships between
mammal hosts and their ectoparasites. Pages 677-723 _i_n_
Ectoparasites of Parana. Field Mus. Nat. Hist. Chicago,
Illinois.

West, R. C. 1956. Mangrove swamps of the Pacific Coast of Columbia.
Ann. Ass. Amer. Geogr. '46: 98-121.

 

 

..
1‘ ‘