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Developments in Primatology:
Progress and Prospects
For other titles published in this series, go to
www.springer.com/series/5852

Anne M. Burrows
● Leanne T. Nash
Editors
The Evolution of
Exudativory in Primates

Editors
Anne M. Burrows
Department of Physical Therapy
Duquesne University
Pittsburgh, PA 15282
and
Department of Anthropology
University of Pittsburgh
Pittsburgh, PA 15260
USA
[email protected]
Leanne T. Nash
School of Human Evolution and Social Change
Arizona State University
Tempe, AZ 85287-2402
USA
[email protected]
ISBN 978-1-4419-6660-5 e-ISBN 978-1-4419-6661-2
DOI 10.1007/978-1-4419-6661-2
Springer New York Dordrecht Heidelberg London
Library of Congress Control Number: 2010936362
© Springer Science+Business Media, LLC 2010
All rights reserved. This work may not be translated or copied in whole or in part without the written
permission of the publisher (Springer Science+Business Media, LLC, 233 Spring Street, New York, NY
10013, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in
connection with any form of information storage and retrieval, electronic adaptation, computer software, or
by similar or dissimilar methodology now known or hereafter developed is forbidden.
The use in this publication of trade names, trademarks, service marks, and similar terms, even if they are
not identified as such, is not to be taken as an expression of opinion as to whether or not they are subject
to proprietary rights.
Cover illustration:
Top image courtesy of K.A.I. Nekaris. A pygmy slow loris, Nycticebus pygmaeus, from Seima
Biodiversity Conservation Area, Cambodia. She was resting during a night of radio tracking, nearby her
parked twins!
Bottom image courtesy of Simon Bearder. The gum is oozing from an Acacia karoo tree used by Galago
moholi as a year round source of energy and minerals, but it is also an important fallback food when
insects are no longer available in the cold, dry winters in South Africa.
Printed on acid-free paper
Springer is part of Springer Science+Business Media (www.springer.com)

Anne dedicates this volume to her family
whose constant support made it possible.
Leanne would like to dedicate this volume to
her parents and siblings, who were always
supportive even when they didn’t fully
understand, and to her spouse, Mike, who
does understand and without whom the work
would not have happened.

vii
I first became involved in research into primate behavior and ecology in 1968, over
40 years ago, driven by a quest for a better understanding of the natural context of
primate evolution. At that time, it was virtually unknown that primates can exploit
exudates as a major food source. I was certainly unaware of this myself. By good
fortune, I was awarded a postdoctoral grant to work on lemurs with Jean-Jacques
Petter in the general ecology division of the Muséum National d’Histoire Naturelle
in Brunoy, France. This provided the launching-pad for my first field study of
lesser mouse lemurs in Madagascar, during which I gained my initial inklings of
exudate feeding. It was also in Brunoy that I met up with Pierre Charles-
Dominique, who introduced me to pioneering observations of exudate feeding he
had made during his field study of five lorisiform species in Gabon. This opened
my eyes to a key feeding adaptation that has now been reported for at least 69
primate species in 12 families (Smith, Chap. 3) – almost 20% of extant primate
species. So exudativory is now firmly established as a dietary category for pri-
mates, alongside the long-recognized classes of faunivory (including insectivory),
frugivory, and folivory.
Soon after I encountered Charles-Dominique, he published the first synthetic
account of his Gabon field study in a French language journal (Charles-Dominique
1971). Convinced by the particular importance of his research, I offered to work on
an English translation, which eventually appeared in book form some 6 years later
(Charles-Dominique 1977). In the process, I learned more about exudativory. For
my own part, I included some preliminary comments on exudate feeding in my
publication on the 1968 study of lesser mouse lemurs (Martin 1972a). In an over-
view of the adaptive radiation of lemurs published in the same year (Martin 1972b),
I made several brief comments on the general significance of exudate feeding. In
particular, I suggested a connection with the tooth-scraper: “Field observations
have shown that the smaller-bodied Cheirogaleinae and Galaginae use the tooth-
scraper to gather plant exudates, and it is likely that the horizontal arrangement of
these anterior teeth has been primarily developed for scraping and prising.”
Although I had thus become aware of the potential significance of exudate feed-
ing, my first really intensive exposure to it came with a 2-year radio-tracking field
study of behavior and ecology of lesser bushbabies in South Africa (1975–1977).
Foreword

viii Foreword
Simon Bearder, the postdoctoral scientist in this investigation who did the lion’s
share of hands-on work in the field, had previously completed an MSc thesis on our
study species, Galago moholi. As a result, he was already quite familiar with exu-
date feeding in this species. I still remember my first night of observations in this
new study. It was a complete revelation to me as Simon led me from one Acacia
tree to another, pointing out sites and traces on the trunks where the bushbabies
came to feed on exudates. Despite the obvious importance of exudates in the diet,
it was not at all easy to see exactly how the bushbabies harvested this key resource.
On rare occasions, it proved possible to observe active use of the tooth-scraper in
exudate feeding. For this, the mouth was held slightly ajar and jerky to-and-fro head
movements were made with the lower jaw applied to the surface of a branch or
trunk. However, the clearest evidence of tooth-scraper use in feeding came from an
incidental, indirect source. In order to fit and remove radio transmitters, we had to
trap our bushbabies regularly. Capture was achieved with large traps placed in trees,
left permanently in situ and regularly supplied with bait poured onto a baseboard.
The bait – containing honey, treacle, peanut butter and banana – was initially a fluid
paste. However, bait left-overs became quite hard when dry. Bushbabies often
entered traps to feed on hardened bait at times when no trapping was conducted.
When the traps were taken down at the end of the 2-year study, all baseboards were
densely covered with characteristic sets of short, parallel scratches that had clearly
been made with the tooth-scraper (Bearder and Martin 1980).
As additional field reports accumulated, notably from a major study of five
sympatric nocturnal lemur species in western Madagascar (Charles-Dominique
et al. 1980), it became clear that exudate feeding was prevalent among small-
bodied, nocturnal strepsirrhine primates (lemurs and lorises). Moreover, it emerged
that fork-crowned lemurs (Phaner) in Madagascar, like needle-clawed bushbabies
(Euoticus) in Africa, are specialist exudate feeders. In parallel to studies on noctur-
nal strepsirrhines, equally striking evidence of exudate feeding was emerging for
small-bodied, diurnal, clawed New World monkeys (Callitrichidae). Napier and
Napier (1967) noted that marmosets (Callithrix, Cebuella) have a “short-tusked
condition,” with relatively long lower incisors and inconspicuous lower canines that
do not project far above the crowns of cheek teeth. By contrast, tamarins (Saguinus,
Leontopithecus) have a “long-tusked condition,” with relatively short incisors and
prominent canines in the lower jaw. This morphological distinction separates mar-
mosets not only from tamarins but also from Goeldi’s monkey (Callimico). It was
later noted (Coimbra-Filho and Mittermeier 1976, 1977) that several marmoset spe-
cies use their lower anterior teeth to perforate tree bark and thus actively stimulate
the flow of exudates. Tamarins and Goeldi’s monkeys have never been observed to
do this, although various species do feed on exudates (Garber and Porter, Chap. 4).
It was also reported that enamel is lacking on the internal (lingual) face of lower
incisors in marmosets (Rosenberger 1978). This resembles the condition seen in
anterior gnawing teeth of lagomorphs, rodents, and Daubentonia. That condition is
regarded as an adaptation for maintenance of a sharp cutting edge. The short-tusked
condition in marmosets was accordingly interpreted as a special adaptation for
actively gouging holes in trees to feed on exudates. New research on enamel prisms

ixForeword
in the anterior lower dentition has revealed that marmosets, but not tamarins, have
clear decussation patterns indicating strengthening to meet the demands of gouging
(Ravosa et al., Chap. 9).
Nearly 25 years ago Nash (1986) effectively reviewed dietary, behavioral, and
morphological correlates of exudativory in primates, bringing together evidence for
both nocturnal strepsirrhines and diurnal callitrichids. Most cases of exudativory
come from relatively small-bodied, essentially arboreal primates, but there are
exceptions. As a rare adaptation among adult Old World monkeys, East African
patas monkeys (Erythrocebus patas) feed primarily on exudates (Isbell 1998).
Exudate feeding has also been reported for yellow baboons, and this is one of the
most important dietary energy sources for juveniles (Altmann 1998). Chimpanzees
have also been reported to feed on exudates, although the daily energy intake from
this source is quite limited (Ushida et al. 2006). Even with the more limited infor-
mation available 25 years ago, Nash’s review was able to establish quite clearly the
importance of exudate feeding for primates.
Accumulating evidence has been accompanied by improving clarity with respect
to basic concepts. In the first place, it is important to distinguish several distinct
kinds of plant exudates that may be consumed by primates: gums, saps, nectar,
latex, and resins (Bearder and Martin 1980; Power, Chap. 2). At the outset, I was
myself confused about these different categories, referring to gums, saps, and resins
interchangeably (Martin, 1972b). However, I soon realized the errors of my ways
(Bearder and Martin 1980). In fact, latex is rarely consumed and resins are shunned,
so the main exudates eaten by primates are gums, saps, and nectar, with gum at the
forefront. Hence, the primary form of exudativory is gum feeding (gummivory). In
this connection, it is also important to distinguish between primates that can gouge
holes in tree trunks and branches (and therefore gain access to saps as well as gums)
and those that either cannot or do not and merely scrape away superficial exudates
(gums). It has long been known that marmosets actually gouge holes, and some kind
of clearly identifiable dental adaptation is therefore to be expected. By contrast, most
strepsirrhines do not gouge but rely on scraping and licking to harvest exudates, so
dental specializations may be correspondingly more subtle.
I originally believed that the tooth-scraper of small-bodied strepsirrhines such as
Microcebus and Galago is too fragile to permit actual gouging. However, apical
wear on the tooth-scraper is seen in relatively old individuals of Galago moholi
(Bearder and Martin 1980), and gouging of some kind has been reported for Phaner
(Petter et al. 1971). But there is now a convincing field report, based on close-up
observations, that Microcebus griseorufus – a specialist gum-feeder among lesser
mouse lemurs – definitely uses its tooth-scraper to stimulate gum flow (Génin
et al., Chap. 6). In another direction, some years ago, I was quite taken by surprise
by an incidental observation made at the Psychological Institute of the University
of Zürich. Gustl Anzenberger, who managed a primate breeding colony, had pro-
vided housing for a pair of pygmy slow lorises (Nycticebus pygmaeus). One day, he
told me that I should come and look at something that would surely interest me. On
various wooden fittings taken from the cage, including both branches and plywood
panels, deep pits were clearly recognizable. Gustl had unmistakably observed the

x Foreword
pygmy slow lorises gouging these pits with their tooth-scrapers. I therefore confi-
dently expected that studies of Nycticebus pygmaeus would reveal exudativory to
be a prominent part of its feeding behavior. I had to wait a few years, but eventually
combined circumstantial evidence from a field survey in Vietnam and from obser-
vations in captivity (at Duke University Primate Center) did indicate that Nycticebus
pygmaeus may be a specialized gummivore (Tan and Drake 2001). It was explicitly
suggested that pygmy slow lorises use the tooth-scraper to chisel away the cam-
bium layer in search of exudates. In fact, it has since been reported that larger-
bodied slow lorises (Nycticebus coucang) also exhibit gouging behavior. Field
observations in Sumatra revealed that slow lorises perforate the superficial layer of
the cambium of trees or lianas with their tooth-scraper (Nekaris et al., Chap. 8). The
onset of gouging is so loud that it is audible at a distance of 30 feet and can be used
as a means of locating individuals at night.
It is also important to distinguish between obligate (“specialist”) exudativores
and facultative (“nonspecialist”) consumers (Nash and Burrows, Chap. 1). Examples
are Euoticus in comparison to other galagids, Phaner as contrasted with most other
cheirogaleids, and marmosets as opposed to other callitrichids. As a rule, gouging
is restricted to (but not universal among) obligate consumers for which exudates
typically make up a large part of the diet. By contrast, facultative consumers depend
only to a relatively small extent on gums and never gouge. So their feeding on
exudates is necessarily confined to gums. It is a moot point whether Microcebus
griseorufus (whose diet includes more than 75% exudates) is really an obligate
exudativore or merely a typical lesser mouse lemur that happens to occupy a habitat
where predominant gum feeding is the only option. But there is a clear expectation
that Nycticebus species, which clearly gouge, will be found to be committed exu-
dativores when appropriate long-term field studies have been conducted.
The fact remains that most gum-eating strepsirrhines do not gouge trees to
obtain sap. Instead, they rely on harvesting superficial gum deposits that are pro-
duced by trees in response to damage. In our field study of Galago moholi, for
example, this was true of the Acacia trees that provided the main source of gum. It
emerged that wood-eating (xylophagous) larvae of various insects bored channels
beneath the tree surface: long-horned beetles (Cerambycidae), jewel beetles
(Buprestidae), click beetles (Elateridae) and carpenter moths (family Cossidae).
Gum was then liberated through surface apertures made by the boring insects, par-
ticularly when they eventually emerged from the host tree (Bearder and Martin
1980). Similar observations have been reported for gum-flows in Madagascar.
Larvae of long-horned beetles and click beetles reportedly chew tunnels in trunks
of Alantsilodendron trees that serve as the main source of gum (Génin et al.,
Chap. 6, this volume). However, I must admit that I simply do not understand what
is going on with respect to the trees that serve as gum sources. Exudation of gum
is generally seen as a tree’s response to damage, sometimes caused by wood-boring
insects, but sometimes resulting from breakage of a branch. Yet it is not at all clear
why only certain trees produce gum, why they sometimes produce large quantities
over an extended period of time, nor why most gums are edible and quite nutritious
rather than laced with toxins. Nash (1989) showed experimentally that addition of

xiForeword
tannins to gum reduces acceptability. The relationship between gum-producing
trees, wood-boring insect larvae, and exudativorous primates is something that
surely deserves more attention in future research.
Gums have one distinct advantage as a food source in that they generally seem
to be available throughout the year, with no marked seasonal pattern of variation.
Some primates, such as Galago moholi, use them as a year-round food source
(Bearder and Martin 1980), whereas others consume them only seasonally, as in
the case of Microcebus murinus (Joly-Radko and Zimmermann, Chap. 7). Overall,
it seems that gums are particularly important during the dry season, when fruits
are often relatively scarce. For this reason, they are often seen as a fall-back food
resource. However, Génin et al. (Chap. 6) suggest the alternative hypothesis that
gummivory is typical of hypervariable environments influenced by El Niño-related
droughts. But in any event dry conditions seem to provide the key. Provided that
they can be digested, gums provide a rich source of carbohydrates (Power,
Chap. 2). Most gummivorous primates have an enlarged caecum, housing symbi-
otic bacteria that can digest the gums. Yet observations of Microcebus griseorufus
indicate that gummivory in this species may not involve a major digestive chal-
lenge, so further study is needed (Génin et al., Chap. 6). Analysis of Acacia gums
gives the misleading impression that exudate composition is fairly consistent.
However, study of gums from other genera has revealed that there is in fact con-
siderable variability. For instance gums of Alantsilodendron in Madagascar are
rich in proteins, whereas protein is only present as a trace component in gums of
Acacia. Minerals such as calcium represent another potentially important compo-
nent of exudates (Garber 1984), although there is no direct evidence that they are
nutritionally important for that reason. One early suggestion was that the high
calcium: phosphorus ratio in Acacia gums may complement the reversed ratio
found in insects and some fruits especially during gestation and lactation (Bearder
and Martin 1980). Ushida et al. (2006) estimated that the average daily intake of
Albizia gum by chimpanzees could meet the entire daily requirement of calcium
and several other minerals, despite the fact that gum makes up a relatively small
part of the diet (Power, Chap. 2).
Exudativory is obviously an important feeding adaptation for various strepsir-
rhines (several cheirogeleids, galagids, and lorisids), callitrichids, and certain
Old World monkeys and apes. Specialization on exudativory developed conver-
gently at least three times during primate evolution and probably more often,
particularly if plesiadapiforms are included (Rosenberger, Chap. 14). Accordingly,
the adaptations associated with gouging and or scraping may differ quite mark-
edly among taxa, notably between strepsirrhines and callitrichids. Yet there are
some general similarities in skull form and jaw mechanics (Vinyard et al. 2003;
Ravosa et al., Chap. 9; Mork et al., Chap. 10). Adaptation for a wide gape gener-
ally seems to be important for gum feeding, reflected by a relatively low-slung
jaw joint with antero-posterior elongation of articular surfaces. On the other
hand, the balance of evidence has discounted an initial expectation that gouging,
if not scraping, would generate larger bite forces requiring special adaptations of
the skull and jaws.

xii Foreword
One long-term goal is a broad evolutionary perspective on exudativory. In her
1986 review Nash aptly stated: “Understanding the biological bases of gummivory
will be of value in interpreting the anatomy and modeling the behavior of early
primates.” One key question that arises here is whether there is an evolutionary
connection between exudativory and the tooth-scraper of strepsirrhine primates. The
six-tooth scraper incorporating the canines and incisors in the lower jaw is almost
certainly a shared derived feature of strepsirrhines that was present in their last
common ancestor. It has long been held that the tooth-scraper evolved in specific
connection with grooming behavior, living up to the alternative name “toothcomb”
(Rosenberger and Strasser 1985; Rosenberger, Chap. 14). However, my own pre-
ferred hypothesis is that evolution of the strepsirrhine tooth-scraper was primarily
connected with exudativory (Martin 1972b, 1979). Of course, it is quite evident that
extant strepsirrhines do generally use their tooth-scrapers in grooming. And there is
also abundant evidence, briefly reviewed above, that numerous strepsirrhines
actively use the tooth-scraper when feeding on exudates. It is also generally accepted
that the specialized gummivores Euoticus and Phaner have distinctive tooth-scrap-
ers. However, the association between tooth-scraper dimensions and exudativory is
not particularly strong when seen across strepsirrhines generally (Eaglen 1986).
Moreover, comparison of specialized exudativores, moderate exudativores and non-
exudativores among galagids provides only limited support for a connection between
tooth-scraper dimensions and exudativory (Burrows and Nash, Chap. 11). But it
must also be recognized that, other than the observation that modern strepsirrhines
do generally use the lower anterior teeth in grooming, there is also no confirmatory
evidence linking grooming to the tooth-scraper. Lots of mammals use whatever
anterior teeth they have for grooming. Examination of hair length in strepsirrhines
showed no relationship with dimensions of the tooth-scraper (Martin 1979). Indeed,
the committed exudativores Euoticus and Phaner both have relatively short hair in
comparison with other strepsirrhines.
I see two basic problems with the hypothesis that the strepsirrhine toothcomb
evolved exclusively for grooming: First, mammalian teeth are generally connected with
feeding behavior, and persuasive arguments are necessary to support evolution in a
nonfeeding context. Second, nobody has ever suggested why strepsirrhines should have
needed a special dental adaptation for grooming. This links up with the fundamental
problem that the tooth-scraper of strepsirrhine primates (including the lower canines) is
unique among mammals, so we have no parallel cases to test hypotheses regarding
grooming or feeding. This issue remains unresolved; the controversy continues.
In closing, I will take the liberty of embarking on a flight of fancy. Bear with me
and accept, for the sake of argument, that the tooth-scraper emerged in ancestral
strepsirrhines in association with scraping (but not gouging) as a means of harvest-
ing exudates. The tooth-scraper was doubtless used for grooming as well, but that
does not affect my argument. Exudativory is essentially an arboreal behavior, so it
would be logical for it to develop in early primates. Now let us consider the strange case
of the aye-aye (Daubentonia). The aye-aye no longer has a typical strepsirrhine
tooth-scraper containing six teeth. Instead, it has a continuously growing, chisel-
like incisor on either side of the lower jaw. The condition in Daubentonia was

xiiiForeword
undoubtedly derived from the original strepsirrhine tooth-scraper by loss of teeth
and other modifications. The chisel-like incisors and filiform, highly mobile middle
finger of the aye-aye represent adaptations for finding and consuming wood-boring
larvae in the trunks of trees. These are the very same larvae that provoke production
of exudates by the host tree. So here’s the thing: Maybe original use of the tooth-
comb to scrape away exudates gradually led to longer and stronger incisors, pene-
tration of bark and eventual feeding on the wood-boring larvae themselves.
Molecular evidence has now convincingly demonstrated that the aye-aye branched
away at the base of the lemuriform radiation (Pastorini et al. 2003; Horvath et al.
2008). So evolution of the condition in Daubentonia according to the scenario just
presented would require an ancestral condition with a six-tooth scraper associated
with exudativory. It has long been accepted that the striped possum of Australasia
(Dactylopsila) provides a marsupial analog to the aye-aye (Cartmill 1974).
Interestingly, Dactylopsila belongs to the same family (Petauridae) as the sugar
glider (Petaurus breviceps), an Australian marsupial that feeds on exudates of
Acacia trees (Smith 1982). So perhaps the evolution of Daubentonia from an ances-
tral, exudativorous strepsirrhine is also paralleled by the evolution of Dactylopsila
from an ancestral, exudativorous marsupial.
Acknowledgments First and foremost, I would like to thank the editors of this volume – Anne
Burrows and Leanne Nash – for kindly inviting me to write this foreword. The chapters grew out
of their well-organized and highly informative symposium on exudativory, held at the 2008
Congress of the International Primatological Society (IPS) in Edinburgh, Scotland. Having par-
ticipated in the early beginnings of research into exudativory, I made certain that attendance at
that symposium was one of my top priorities. I was delighted by the quality of the presentations.
It was also a pleasure to see how many advances had been made in our understanding of an
important primate feeding category that was virtually unrecognized 40 years ago. I learned a
great deal, both from presentations at the symposium and from the more detailed information
provided in the chapters contained in this volume. My pleasure was further heightened when
Anne and Leanne invited me to write a foreword straight after the symposium. I feel honoured
to be involved in this way, as The Evolution of Exudativory in Primates undoubtedly represents
a watershed in our understanding of this fascinating topic. The 14 chapters effectively review
current information from a wide array of disciplines: behavior, ecology, nutrition, primate evolu-
tion, morphology, and conservation. They also provide pointers to future research that will hope-
fully answer several puzzling questions that remain open. Let me conclude by acknowledging the
great debt that we owe to the dedicated primate fieldworkers who discovered the phenomenon of
exudativory and have generated a continuing flow of vital information from natural habitats.
Robert D. Martin
The Field Museum, Chicago, IL USA
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xiv Foreword
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monkey (Erythrocebus patas pyrrhonotus). Am J Primatol 45:381–398
13. Martin RD (1972a) A preliminary field-study of the lesser mouse lemur (Microcebus murinus,
J.F. Miller 1777). Z Tierpsychol Suppl 9:43–89
14. Martin RD (1972b) Adaptive radiation and behaviour of the Malagasy lemurs. Phil Trans Roy
Soc Lond B 264:295–352
15. Martin RD (1979) Phylogenetic aspects of prosimian behavior. In The study of prosimian
behavior (eds Doyle GA, Martin RD). Academic Press, New York
16. Napier JR, Napier PH (1967) A handbook of living primates. Academic Press, London
17. Nash LT (1986) Dietary, behavioral, and morphological aspects of gummivory in primates.
Yrbk Phys Anthropol 29:113–137
18. Nash LT (1989) Galagos and gummivory. Hum Evol 4:199–206
19. Pastorini J, Thalmann U, Martin RD (2003) A molecular approach to comparative phylogeog-
raphy of extant Malagasy lemurs. Proc Natl Acad Sci USA 100:5879–5884
20. Petter J-J, Schilling A, Pariente G (1971) Observations éco-éthologiques sur deux lémuriens
malgaches nocturnes: Phaner furcifer et Microcebus coquereli. Terre Vie 118:287–327.
21. Rosenberger AL, Strasser ME (1985) Toothcomb origins: support for the grooming hypoth-
esis. Primates 26:76–85
22. Rosenberger AL (1978) Loss of incisor enamel in marmosets. J Mammal 59:207–208
23. Smith AP (1982) Diet and feeding strategies of the marsupial sugar glider in temperate
Australia. J Anim Ecol 51:149–166
24. Tan CL, Drake JH (2001) Evidence of tree gouging and exudate eating in pygmy slow lorises
(Nycticebus pygmaeus). Folia Primatol 72:37–39
25. Ushida K, Fujita S, Ohasgi G (2006) Nutritional significance of the selective ingestion of
Albizia zygia gum exudate by wild chimpanzees in Bossou, Guinea. Am J Primatol
68:143–151
26. Vinyard CJ, Wall CE, Williams SH, Hylander WL (2003) Comparative functional analysis of
skull morphology of tree-gouging primates. Am J Phys Anthropol 120:153–170

xv
Acknowledgments
This volume is a product of a symposium entitled “The evolution of exudativory in
primates” held at the 22nd Congress of the International Primatological Society in
Edinburgh, Scotland in 2008. We would like to offer our sincere thanks to the
numerous people who assisted this edited volume throughout its various stages. The
reviewers provided their expert insights and commentary. Melissa Higgs at Springer
was always enthusiastic and unfailingly guided and encouraged us. Bob Martin not
only provided a wonderful Foreword, but inspired much of the work represented
here starting over three decades ago. Our most special thanks are due to the con-
tributors whose steadfast labors and patience helped bring this volume to its ulti-
mate materialization.

xvii
Contents
1 Introduction: Advances and Remaining Sticky Issues
in the Understanding of Exudativory in Primates. ................................. 1
Leanne T. Nash and Anne M. Burrows
2 Nutritional and Digestive Challenges
to Being a Gum-Feeding Primate. ............................................................ 25
Michael L. Power
3 Exudativory in Primates: Interspecific Patterns. .................................... 45
Andrew C. Smith
4 The Ecology of Exudate Production and Exudate Feeding
in Saguinus and Callimico. ........................................................................ 89
Paul A. Garber and Leila M. Porter
5 Influences on Gum Feeding in Primates. ................................................. 109
Andrew C. Smith
6 Gummivory in Cheirogaleids: Primitive Retention
or Adaptation to Hypervariable Environments?. ................................... 123
Fabian G.S. Génin, Judith C. Masters, and Jorg U. Ganzhorn
7 Seasonality in Gum and Honeydew Feeding
in Gray Mouse Lemurs. ............................................................................. 141
Marine Joly-Radko and Elke Zimmermann
8 Comparative Ecology of Exudate Feeding by Lorises
(Nycticebus, Loris) and Pottos (Perodicticus, Arctocebus). ...................... 155
K. Anne-Isola Nekaris, Carly R. Starr, Rebecca L. Collins
and Angelina Wilson

xviii Contents
9 Exudativory and Primate Skull Form. ................................................... 169
Matthew J. Ravosa, Russell T. Hogg, and Christopher J. Vinyard
10 A Comparative Analysis of the Articular Cartilage
in the Temporomandibular Joint of Gouging
and Nongouging New World Monkeys. .................................................. 187
Amy L. Mork, Walter E. Horton, and Christopher J. Vinyard
11 Searching for Dental Signals of Exudativory in Galagos. .................... 211
Anne M. Burrows and Leanne T. Nash
12 A Guide to Galago Diversity: Getting a Grip
on How Best to Chew Gum. .................................................................... 235
Isobel R. Stephenson, Simon K. Bearder, Guiseppe Donati,
and Johann Karlsson
13 Tongue Morphology in Infant
and Adult Bushbabies (Otolemur spp.). ................................................. 257
Beth A. Docherty, Laura J. Alport, Kunwar P. Bhatnagar,
Anne M. Burrows, and Timothy D. Smith
14 Adaptive Profile Versus Adaptive Specialization:
Fossils and Gummivory in Early Primate Evolution. ........................... 273
Alfred L. Rosenberger
Index. ................................................................................................................. 297

xix
Contributors
Laura J. Alport
Department of Anthropology, University of Texas at Austin, Austin,
TX 78712, USA
Simon K. Bearder
Nocturnal Primate Research Group, Department of Anthropology and Geography,
School of Social Sciences and Law, Oxford Brookes University,
Oxford,OX3 0BP, UK
Kunwar P. Bhatnagar
Department of Anatomical Sciences and Neurobiology, University of Louisville
School of Medicine, Louisville, KY 40292, USA
Anne M. Burrows
Department of Physical Therapy, Duquesne University, Pittsburgh, PA 15282,
USA, Department of Anthropology, University of Pittsburgh, Pittsburgh,
PA 15260, USA
Rebecca L. Collins
Nocturnal Primate Research Group, Department of Anthropology and Geography,
School of Social Sciences and Law, Oxford Brookes University,
Oxford OX3 0BP, UK
Beth A. Docherty
Department of Physical Therapy, Duquesne University, Pittsburgh,
PA 15282, USA
Guiseppe Donati
Nocturnal Primate Research Group, Department of Anthropology and Geography,
School of Social Sciences and Law, Oxford Brookes University,
Oxford OX3 0BP, UK
Jorg U. Ganzhorn
Department of Zoology, University of Fort Hare, Private Bag X1314,
Alice 5700, South Africa

xx Contributors
Paul A. Garber
Department of Anthropology, University of Illinois, 109, Davenport Hall,
607 S Mathews Ave, Urbana, IL 61801, USA
Fabien G.S. Génin
Department of Zoology, University of Fort Hare, Private Bag X1314,
Alice 5700, South Africa
Russell T. Hogg
Department of Pathology and Anatomical Sciences, School of Medicine,
University of Missouri, Columbia, MO 65212, USA
Walter E. Horton, Jr.
Department of Anatomy and Neurobiology, Northeastern Ohio Universities
College of Medicine (NEOUCOM), Rootstown, OH 44272, USA
Marine Joly-Radko
Institut fuer Zoologie, Tieraerztliche Hochschule Hannover, Buenteweg 17,
Hannover 30559, Germany
Johann Karlsson
Nocturnal Primate Research Group, Department of Anthropology and Geography,
School of Social Sciences and Law, Oxford Brookes University,
Oxford OX3 0BP, UK
Judith C. Masters
Department of Zoology, University of Fort Hare, Private Bag X1314, Alice 5700,
South Africa
Amy L. Mork
Department of Anatomy and Neurobiology, Northeastern Ohio Universities
College of Medicine (NEOUCOM), Rootstown, OH 44272, USA
Leanne T. Nash
School of Human Evolution and Social Change, Arizona State University,
Tempe, AZ 85287-2402, USA
Angelina Wilson
TRAFFIC International, 219a Huntingdon Road Cambridge, CB3 0DL, UK
K. Anne-Isola Nekaris
Nocturnal Primate Research Group, Department of Anthropology and Geography,
School of Social Sciences and Law, Oxford Brookes University,
Oxford OX3 0BP, UK
Leila M. Porter
Department of Anthropology, University of Illinois, 109, Davenport Hall,
607s Mathews Ave, Urbana, IL 61801, USA

xxiContributors
Michael L. Power
Nutrition Laboratory, Smithsonian Conservation Biology Institute, National
Zoological Park, P.O. Box 37012, MRC 5503, Washington, DC 20013-7012, USA
Research Department, American College of Obstetricians and Gynecologists,
Washington, DC 20024, USA
Matthew J. Ravosa
Department of Pathology and Anatomical Sciences, School of Medicine,
University of Missouri, Columbia, MO 65212, USA; Division of Mammals,
Department of Zoology, Field Museum of Natural History, Chicago,
IL 60605-2496, USA
Alfred L. Rosenberger
Department of Anthropology and Archaeology, Brooklyn College,
The City University of New York, Brooklyn, NY 11210, USA;
The Graduate Center, The City University of New York, New York, NY, USA
New York Consortium in Primatology (NYCEP), NY, USA
and
Department of Mammalogy, The American Museum of Natural History,
New York, NY 10024-5192, USA
Andrew C. Smith
Animal and Environmental Research Group, Department of Life Sciences,
Anglia Ruskin University, East Road, CB1 1PT Cambridge, UK
Timothy D. Smith
School of Physical Therapy, Slippery Rock University, Slippery Rock, PA 16057,
USA; Department of Anthropology, 3302 WWPH, University of Pittsburgh,
Pittsburgh, PA 15260, USA
Carly R. Starr
School of Animal Studies, University of Queensland, Gatton, Queensland 4072,
Australia
Isobel R. Stephenson
Department of Anthropology and Geography, School of Social Sciences and Law,
Oxford Brookes University, Gipsy Lane, Oxford, OX3 0BP, UK
Christopher J. Vinyard
Department of Anatomy and Neurobiology, Northeastern Ohio Universities
College of Medicine (NEOUCOM), Rootstown OH 44272, USA
Elke Zimmermann
Institute of Zoology, University of Veterinary Medicine Hanover,
Hanover, Germany

1A.M. Burrows and L.T. Nash (eds.), The Evolution of Exudativory in Primates,
Developments in Primatology: Progress and Prospects,
DOI 10.1007/978-1-4419-6661-2_1, © Springer Science+Business Media, LLC 2010
Abstract In the 25 years since the last synthesis on this topic was published, there
has been a marked increase in the appreciation of exudativory as a primate dietary
strategy and investigations of its morphological correlates that appear to be adaptations
to exudates as food. At least 75 species of primates consume some exudates. Variability
of diet among marmosets and tamarins precludes simple classifications of the former
as year-round specialists vs. the latter as always facultative seasonal users of exudates.
Differences in exudate use among callithrichines, now also including callimico as an
exudativore, are associated with apparent adaptations in gut anatomy and functioning,
a suite of dental and jaw features, and some features of socioecology and life-history.
Among strepsirrhines, several Nycticebus species are newly known to gouge to eat
gum, variability among mouse lemurs in gum use has been documented, but little added
work has improved our knowledge of variation in exudate use in galagos. For these
taxa, much less is understood about possible morphological, behavioral and life-history
adaptations and detailed descriptions of behaviors associated with exudate acquisition
are needed from the field. The ability to identify anatomical features that will clarify the
role of exudates in the diets of fossil primates remains a major challenge.
Introduction
The present book, for which this chapter serves as an Introduction, grew out of a
symposium held at the 2008 Congress of the International Primatological Society
(IPS) in Edinburgh, Scotland. This symposium was organized to bring together in
one place and at the same time researchers from many diverse fields (ecology,
behavior, morphology, nutrition, and conservation) that all converged on the topic
L.T. Nash (*)
School of Human Evolution and Social Change, Arizona State University,
Tempe, AZ 85287-2402, USA
e-mail: [email protected]
Chapter 1
Introduction: Advances and Remaining
Sticky Issues in the Understanding
of Exudativory in Primates
Leanne T. Nash and Anne M. Burrows

2 L.T. Nash and A.M. Burrows
of primate exudativory and how it has evolved. Despite the fact that one of us (Nash 1986)
published a review nearly 25 years ago on the dietary, behavioral, and morphological
correlates of primate exudativory, much remains to be understood about this rela-
tively rare dietary niche.
We have envisioned this chapter as both an introduction to the other chapters in
this book and as a brief overview and update of the 1986 review (Nash 1986).
We have blatantly borrowed the overall structure of that review for this chapter.
That review described “aspects” of “gummivory” in primates. The title we have
chosen for this volume is somewhat more ambitious, as our aim is to address the
“evolution of exudativory.” In 1986, exudativory as a dietary niche was relatively
unknown and thoroughly under-studied. Many new developments have occurred
since then including which primates consume gum, the socioecological correlates
of exudativory, behavioral and morphological adaptations to locating, accessing,
consuming, and digesting exudates, and even the role exudate-feeding may have
played in the origin of primates. The encouraging point is that today one paper,
such as this one, cannot be an exhaustive review of the current state of the work on
exudativory in primates. This book approaches that. This increase in appreciation
of the ramifications of exudate-feeding in primate biology is dramatic; however,
so too are the remaining gaps in our knowledge on the topic. Our goal for this
chapter is to provide the context of the remaining chapters. Our greatest goal for
this book is to stimulate further work that lets us get unstuck from some of the
questions that remain.
We have chosen the term “exudativory” in preference to “gummivory” as a
more inclusive term. We have left it to each author to be more specific as the con-
text of their data and discussions dictates. Exudates consumed by primates include
gums, saps, nectar, and, more rarely resins and latex (Power, Chap. 2). Nectar is not
explicitly addressed in this volume, though a number of species such as Eulemur
spp., Galago senegalensis, and Otolemur crassicaudatus will visit flowers and lick
them without destroying them (Sussman and Raven 1978; Charles-Dominique and
Bearder 1979; Nash, pers. obs.). Resins are not eaten by primates but some primates
may consume latex. Latex is sticky, chemically complex, contains proteins and a
variety of secondary compounds, and has evolved as a defense against insect dam-
age to plants in leaves and reproductive parts (e.g., latex in figs) (Agrawal and
Konno 2009). For example, two of the three most frequently eaten plants that
Lepilemur leucopus consumes in southwestern Madagascar at Beza Mahafaly
Reserve have sticky, milky, sometimes irritating, exudate that appears to be latex
(Nash 1998). The chemical composition of such plants remains to be fully explored
(Power, Chap. 2).
The majority of the exudates to be discussed in this book are gums (water-soluble,
viscous exudates found just deep to the bark) and saps (water-soluble, viscous
exudates found deeper in the xylem and phloem). Most of the gums come from
nonreproductive parts of the tree, are produced in response to insect or mechanical
damage, and are not produced to attract animals to service the plant, in contrast to
nectar (Power, Chap. 2). However, pod gums, as consumed by some neotropical
primates (Power, Chap. 2, Smith, Chaps. 3 and 5; Garber and Porter, Chap. 4) may

31 Introduction: Advances and Remaining Sticky Issues
be exceptions to this generalization. Saps present a special problem: in order to be
accessed, the bark of the tree must be broken through sufficiently to reach this
deeply located resource. For the most part, only primates that can specifically
gouge into the bark, such as marmosets, access sap. This leads to an important
distinction between primates which can properly gouge and those which cannot or
do not – a key topic in many of the chapters of this book (see below). However,
Joly-Radko and Zimmermann (Chap. 7) have documented a case of “sap eating by
proxy” in cheirogalieds that eat the “honeydew” produced by sap-sucking insects.
To date, this form of exudativory has not received much attention and involves a
potentially complex web of ecological relationships among primates, insects and
plants that deserves further study.
Such a complex web of ecological interactions has been described among a
gum-eating Australian marsupial, the sugar glider (Petaurus breviceps), wattle spe-
cies (Australian Acacia trees) that produce gum, sheep, beetles, and eucalyptus
trees (Smith 1992). Sugar gliders are convergent with Galago moholi and G. sen-
egalensis in their habitat preference and dependence on Acacia gum in the cold
season when insects are rare (Harcourt 1986). While galagos leap, the small mar-
supials glide between trees. Sugar glider density is positively associated with wattle
(Acacia mearnsii) density (Smith 1982; Suckling 1984). In areas with heavy sheep
grazing, the seedling wattles are grazed so they regenerate poorly. Like G. moholi
and G. senegalensis, gliders incorporate insects as a major portion of their diet in
seasons when they are available. The gliders are an important biological control on
leaf-defoliating scarab beetles. When weather conditions are suitable, beetles prolif-
erate and may defoliate eucalyptus timber species producing a “dieback” condition
that can kill the trees. Thus, key elements for maintaining forest health and dieback
resistance in these rural ecosystems are sugar gliders and wattles. Keeping these
species in the habitat requires controlling grazing to allow wattles to regenerate so
gliders have a winter food supply of gum. To our knowledge, no primate exudati-
vore has been shown to have such a critical role within an ecosystem, but this is due
to our ignorance of their ecological webs. Ideally, we hope this book can stimulate
more research on all types of exudates which will integrate information on the
behavior of exudate eating, the reasons plants or other sources produce exudates,
the composition and distribution (in time and in space) of exudates that primates
eat, the ecological roles of exudates and exudate consumers, and the morphological
adaptations that allow animals to access this food resource.
Returning to the title of this book, by explicitly incorporating the word “evolution”
in the title, we hoped contributors would address possible behavioral and morpho-
logical adaptations in primates associated with exudate consumption and address
these adaptations in the fossil record. We find there is much remaining to be adequately
tested in identifying such adaptations, especially with regards to the fossil record.
Many of the apparent adaptations may be “clade dependant” and not occur in all
primate exudativores. A major issue (see below) is whether there are, as yet, any
reliable signals from the hard anatomy (i.e., skeletal and dental characters) identifiable
in fossils that would truly allow us to document the history of exudate consumption
in the course of primate evolution.

4 L.T. Nash and A.M. Burrows
A note on taxonomic usage is in order. In this chapter, we follow Groves
(2001, 2005) using “strepsirrhines,” “haplorrhines” and “callithrichines.” We have not
attempted to force a single taxonomic usage on all authors. Instead, we have
requested that each provide citations to their choice of taxonomy for the primates.
However, the main differences across chapters are in lower level taxonomy
(e.g., marmosets and tamarins as a separate family or subfamily, appropriate genus
and species names). Most authors follow the same subordinal taxonomy within
primates as used in this chapter.
Exudates as a Primate Dietary Component
Updates on Primate Exudativores
The knowledge of the major genera and higher taxa of primates that consume
considerable amounts of gum has changed over the years and good major over-
views can be found in recent reviews about primates (Committee on Animal
Nutrition 2003; Campbell et al. 2007) and in Smith (Chap. 3). This chapter reviews
over 130 sources to document 69 primates that consume exudates from over 300 plant
species. As was previously known, strepsirrhines and callithrichines consume the
greatest volumes of exudates relative to their entire diet while patas monkeys, some
vervets, and baboons also consume considerable volumes. Occasional use of pod
exudates in Lagothrix and occasional gum consumption in some langurs, macaques,
forest guenons, and chimpanzees are now also known. As Smith points out, his list
likely misses a number of noncallithrichines and non-Malagasy strepsirrhines due
to a lack of comparable detailed field work on such species. It is notable that the
Committee on Animal Nutrition of the National Academy of Sciences has now
recognized “gums” as a major component of some primates’ diets, as this has
implications for captive husbandry (see below and http://www.nap.edu/catalog.
php?record_id=9826#toc).
Among strepsirrhines, one of the most remarkable and newly documented
findings is the use of exudates by a number of Asian lorises which gouge or scrape
to acquire them (Tan and Drake 2001). Nekaris et al. (Chap. 8) amplify these observa-
tions extensively and show that Nycticebus consumes gum. Ironically, though gala-
gos were among the first primates to be noted as major exudativores, there has been
little recent detailed fieldwork on them or any African lorisoids that has focused on
feeding patterns. As some of our earliest information on primate exudativory came
from galagos (Bearder and Martin 1980a) it is ironic that the African strepsirrhines
are now one of the taxa where the least amount of progress has been made in
understanding their diversity of exudate use (Bearder et al. 1995; Pimley et al.
2003, 2005a, b; Nekaris and Bearder 2007), even as the number of species recog-
nized has exploded (Grubb et al. 2003). Thus, it is unknown how these galagos and
pottos compare to Asian lorises in their consumption of exudates and why they differ.

51 Introduction: Advances and Remaining Sticky Issues
We greatly need more detailed information on the diversity of which exudates are
eaten and how they are acquired for both African and Asian strepsirrhines.
Génin has provided important novel data on gummivory in mouse lemurs in western
and southern Madagascar (Génin 2003; Génin et al. 2005; Génin 2007, 2008). For
example, he has shown that gray mouse lemurs compete intraspecifically for access
to gum and females dominate males in accessing gum sources. He also documents
the first case of Phaner dominating Microcebus murinus at gum sources in Kirindy.
In western forests, mouse lemur species eat gum more seasonally or not at all. In
contrast, at Berenty Microcebus griseorufus use gum year round and consume fruit
seasonally, though less gum is used in wet than dry years (Génin 2008; Génin
et al., Chap. 6). They argue that gum may be a more reliable and rapidly renewing
carbohydrate source than fruit where rainfall is unpredictable and argue that gum use
may be a primitive adaptation in all cheirogalieds. They hypothesize that El Niño-
associated unpredictable droughts are associated with the localities around the world
where exudativory is particularly prominent in primates and other rare mammals that
consume gums. The possible diversity among mouse lemur species and populations
in consumption of gum, as with galagos, may provide opportunities for future comparative
work in both behavioral and morphological adaptations to exudativory. As illustrated
by Génin’s work, such comparisons allow for testing the socioecological model as
applied to the ecology and behavior of a relatively nongregarious primate and may
help understand the origins of primate gregariousness.
The hypothesis that the evolution of gummivory is associated with an unpredictable
environment is bolstered by the recent first fieldwork on Allocebus trichotis in
Madagascar which suggests that, as predicted from its dental morphology, gum is
probably a major part of its diet (Biebouw 2009). It is hoped that more detailed
information on its feeding ecology will become available in the near future. As men-
tioned above, more detailed work on consumption of “honeydew” (a sap derivative)
by Cheirogaleus provides another variant on exudate use for comparative work
(Joly-Radko and Zimmermann, Chap. 7).
Turning to haplorrhines, it was recognized in the 1980s that baboons were one
of the largest-bodied primates that eat gum. What is new is the case that Altmann
(1998) makes for the importance of gum in the diet of juvenile baboons. It is a very
important source of energy in the juvenile’s diet, which, in turn, is one of the strongest
correlates of adult female’s fitness. Some populations of chimpanzees consume
gums as part of their diets (Ushida et al. 2006). While it seems that these chimpanzees
gain negligible energy from the gums, they do seem to gain sufficient amounts of
various minerals (calcium, magnesium, manganese, and potassium) to fulfill their
daily requirements for these minerals. Body size, as well as the nature of other
dietary components, may influence different nutritional advantages for exudativory
across primates (Power, Chap. 2).
Isbell’s work comparing gum use in sympatric vervets and patas monkeys is a
model of the possibilities of comparative work in understanding the processual
events in the evolution of exudativory as a dietary niche (Isbell 1998; Isbell et al.
1998). Patas monkeys in East Africa feed primarily on gums, a rarity among adult
Old World monkeys. Isbell and colleagues have shown that the development of this

6 L.T. Nash and A.M. Burrows
odd dietary niche for a relatively large-bodied Old World monkey may have been
driven in part by their notably long limbs which allow them to cover a wide geo-
graphic range in a given time to gather gums. In addition, these large monkeys seem
to be able to subsist primarily on exudates because substantial volumes of arthropods
are associated with the gums and consumed along with them. The consumption of
gum by patas monkeys, baboons, and living humans has even been incorporated
into reconstructions of the diet of early hominins (Copeland 2007).
In the Neotropics, an important outcome of continuing fieldwork is that it is now
clear that a simple distinction between marmosets as year-round gum specialists vs.
tamarins as seasonal gum users is oversimplified, though it is a model that is very
influential in morphological studies of exudativory (see below). There is considerable
diversity in the extent to which gums are eaten within marmosets, within tamarins,
and even within the same genus of these groups (Smith, Chap. 3). The extent to
which this diversity within marmosets or within tamarins is incorporated into anatomical
studies is quite variable. However, it has countered the suggestion that nails were
adaptations to trunk exudativory (Sussman and Kinzey 1984) as it now appears that
the ancestral callithrichine probably engaged in a variety of trunk foraging behaviors,
not just gum eating (Garber et al. 1996). Callimico, whose diet was very poorly
known in 1986, was then thought not to eat exudates. As reported by Garber and
Porter (Chap. 4), callimicos are now known to feed extensively on two items
unusual in primate diets: fungi and both trunk and pod exudates (see also Porter and
Garber 2004; Porter 2007; Porter et al. 2009). Garber and Porter elaborate here on
how important various exudates may be for some nongouging callithrichines
throughout all or much of the year.
Exudates as Fallback Foods and Primate Feeding Adaptations
A number of authors in this volume as well as others (e.g., Harrison and Tardif 1994)
draw a distinction between “specialist” or “obligate” exudativores and “facultative”
or “nonspecialist” consumers of exudates. The criteria for obligate exudativores
generally included use of gum across all seasons and the possession of morphological
features associated with gum procurement or digestion including small body size,
sharp or claw-like nails, dental features associated with gouging, and an expanded
gut (especially the cecum) for fermentation (see further on anatomy below).
A consumption distinction that recurs in many chapters in this volume is the
extent to which gum is used by a given species or population as a “preferred food,”
a “fallback food,” or as a “keystone” food – and if gum sources can fill more than
one of these roles. These concepts are invoked in most chapters in this volume and
sometimes are treated as if the concepts of keystone food and fallback food are
interchangeable. However, it is best to keep the notions separate (Marshall and
Wrangham 2007). The concept of “keystone species” refers to one that is essential
to the structural and functional integrity of an ecological community, not just one
species, but those keystone effects can be context dependent and may be seasonal

71 Introduction: Advances and Remaining Sticky Issues
(Peres 2000; Christianou and Ebenman 2005; Collinge et al. 2008). There is much
discussion of how to define and to identify a keystone species in the ecological
literature that is beyond our objectives here (Davic 2003; Christianou and Ebenman
2005; Hodges 2008, Fedor and Vasas 2009; Jordán et al. 2009). To what extent
exudativorous primates or the plants from which they consume exudates are keystone
species, in the sense that sugar gliders may be in the example presented above, is
not know for any primate. Peres (2000) has made the case for one type of exudate,
the pod gums from Parkia trees, as a keystone resource in some Neotropical forests.
It is clear that for many primates exudates may be important fallback food.
Fallback foods are defined as foods eaten when preferred foods are not available
(Marshall and Wrangham 2007) and are expected to be of poorer nutritional content
and/or more difficult to process than preferred foods. These authors argue that fallback
foods, while often consumed seasonally, in some cases may be eaten year round.
Their key to identifying a fallback food is that it be eaten in amounts inversely
related to the availability of preferred foods. This criterion requires (1) data on the
availability of all food types, and (2) identification of preferred foods. Preferred
foods are those eaten in excess of their availability; neither availability (rarity) nor
frequency of consumption alone defines a preferred item. Preferred foods are argued
to primarily drive harvesting adaptations (perception, spatial navigation, cognition,
and locomotion), while fallback foods shape processing adaptations (dental and sup-
porting morphology, gut anatomy and kinetics, body size, tool use). Marshall and
Wrangham (2 007) distinguish within fallback foods staple fallback foods (those that
can seasonally be the sole food eaten) and filler fallback foods (those that are never
the whole diet). Acacia gum would be a staple fallback food for G. moholi (Bearder
and Martin 1980a). Compared to staple fallback foods, filler fallback foods are
predicted to fluctuate less in availability through time, engender less feeding compe-
tition, allow more stable grouping patterns, and be associated with faster life-histories.
Lambert (2 007) emphasizes that identifying feeding adaptations requires establishing
linkages of variables at all scales of organismal biology: individual intragenerational,
individual intergenerational, population and species. The chapters in this volume
mainly focus on the latter two levels of scale and on craniodental morphological
features.
Both the models of Lambert (2007) and Marshall and Wrangham (2007) may be
helpful in the future in organizing approaches to understanding the adaptive
pressures exudativory presents and may be able to be integrated (Constantino and
Wright 2009; Lambert 2009; Marshall et al. 2009). In order to decide if exudates
are fallback foods we need better information on their bioavailability of nutrients
(Power, Chap. 2) and on measures of their availability across the seasons (see below).
We need to decide if they are staple or filler fallback foods. For example, some of
the newly studied lorises eat gum year round, and others have not yet been studied
across all seasons of the year, so the role of gums as fallback foods is as yet unclear
(Nekaris et al., Chap. 8). Lambert (2007, 2009) suggests we focus on both “fallback
foods” and “fallback strategies.”
As useful as these models may be, the situation of exudate-feeding on trunk and
pod gums in Callimico may not be a close fit to either. Porter et al. (2009) and

8 L.T. Nash and A.M. Burrows
Garber and Porter (Chap. 4) question to what extent gums are fallback foods.
They consider evidence that the evolution of the gut’s ability to digest gum may
have been a result of adaptations to another preferred diet item – fungi. Part of their
critique is based on the possibility that pod gums are more digestible (see below for
further discussion of this point). Thus they question to what extent different sorts
of processing and harvesting adaptations are, or are not, associated with fallback
foods. Génin et al. (Chap. 6) also show the complications in the “fallback” categorization
in that they explicitly introduce the notion that environments that are “hypervariable”
over longer time scales than a single year may be key in understanding which species
eat gum and geographic variation in eating this food. Though the time scale is
different, they, like Garber and Porter (Chap. 4) raise the issue that the association
of “fallback foods” with adaptations to harvesting vs. processing is more complex than
suggested in the Marshall and Wrangham model (Marshall and Wrangham 2007).
The cognitive, social and life-history ramifications of gum use have barely been
addressed. Between the obligate exudativorous marmosets and the facultative exu-
dativorous tamarins, where there is clear morphological distinction in the dentition,
exudate usage has been interpreted to have ramifications in behaviors as diverse as
the ability to delay rewards, the degree of sex differences in territorial behavior,
infant carrying, and group stability (Ferrari and Lopes 1989; Harrison and Tardif
1994; Stevens et al. 2005). The extent to which these differences can be generalized
to other primate taxa, for example, lorisoids or lemuroids, remains to be examined.
For example, while Cebuella may tend to focus their territory on one or a few gum
trees in their small territories, this is not the pattern that is found among galagos.
The lack of good field or captive data comparing cognitive or social differences
among galagos and lorises that do and do not use gums extensively would be infor-
mative. It is clear that the general social differences among nocturnal strepsirrhines
do not covary in a simple way with the use of exudates for either lemurs (Schülke
and Ostner 2005) or for galagos and lorises (Bearder and Martin 1980b; Clark 1985;
Harcourt and Nash 1986a; Nash and Harcourt 1986; Harcourt and Bearder 1989;
Nekaris and Bearder 2007). It has been argued that Phaner and some marmosets,
though not Cebuella, share a situation where the use of reliably located gum
resources that are quickly depleted, rapidly renewing, and are monopolizable but
not clumped, allows group members to feed near each other. This sets up a situation
where the balance of within and between group scramble and contest feeding
competition favors a social and life-history pattern of delayed natal dispersal
(Schülke 2003; Schülke and Kappeler 2003) which is absent in sympatric relatives
of these species which do not use gum (i.e., Cheirogaleus or tamarins).
Factors Influencing Selectivity in Exudate-Feeding
Factors influencing selectivity include those involved in both the harvesting and
processing of exudates. We have much to learn about which primates eat exudates,
which plant species are accessed, which exudates are used, when exudates are

91 Introduction: Advances and Remaining Sticky Issues
consumed, and why a particular location or “glob” of exudate is chosen. Globs may
be large or tiny relative to the consumer, old and hard or young and liquid, crystallized
or liquid under a dried “skin,” and variously colored (and presumably varying in
taste and scent). Globs may also be located in different parts of the plant. We are
only beginning to understand the factors that influence why exudates are eaten by
a primate at a particular place and time. Some of the main challenges are (1) under-
standing the nutritional gain from exudates and the problems exudates present to
digestion, (2) identifying costs and benefits to the animal of harvesting different
types of exudates based on where the plant produces them and whether or not
harvesting exudates has any benefits to the plant producer, and (3) from the
perspective of primate consumers, measuring availability of an exudate and “patchiness”
of its distribution in time and space.
Digestive Challenges and Nutrients in Gum
Since most of the exudates consumed by primates are gums, it is not surprising that
most work on the nutritional benefits and digestive problems of exudate consumption
has been directed at gums. As detailed by Power (Chap. 2), gums are water soluble,
low in protein, high in some minerals, and mostly beta-linked complex polysac-
charides which are not digestible by mammalian enzymes. They are assumed to
mainly be eaten for their energy content, but the carbohydrates must be fermented
to be bioavailable. However, both between and within plant species, there are
differences in the solubility (which influences fermentability), constituent sugars,
and secondary compounds (“antinutrients”) found in gums (Génin et al., Chap. 6,
Hladik et al. 1980; Lambert 1998; Génin 2003; Wiens et al. 2006). For example,
we know that exudates from plants of the pea-family (Fabaceae) are favored across
most primates (Smith, Chap. 3) and specifically by Asian lorises (Nekaris et al.,
Chap. 8) but we don’t know why. Lemurs seem to have developed the ability to
tolerate a variety of secondary compounds (Ganzhorn 1992). Do they differ from
other strepsirrhines in selecting exudates based on these antinutrients (Reed and
Bidner 2004)? Power (Chap. 2) also points out that commonly used conversion
factors for protein content, based on nitrogen content, may be problematic and
inflated, and not the same as “bioavailability.” Though relative values within a
study may be correct, this may cause problems comparing across studies of different
primates and different plants.
In addition to energy content, early hypotheses of gum consumption suggested
that since it was high in calcium and low in phosphorus, it complemented the
reversed ratio found in insects and some fruits (Bearder and Martin 1980a), especially
during gestation and lactation (Garber 1984). In a review across primates, Smith
(Chap. 5) does not find support for the “cost of reproduction” hypothesis of gum
consumption. However, Garber and Porter (Chap. 4) argue that seasonal differences
in the nutrient and the anti-nutrient contents of gums may be a factor in gum
consumption.

10 L.T. Nash and A.M. Burrows
Digestible Energy form of Trunk Gum vs. Pod Gum vs. Honeydew
There are various suggestions or direct claims that different types of exudates differ
in the nutrients they provide to primates. Joly-Radko and Zimmermann (Chap. 7)
indicate that when both are available in a mouse lemur’s range, honeydew is
preferred to gum. This may imply that it is more nutritious than gum, since the sap
has been predigested by the insects that exude the honeydew. Based on the notion
that plants make their parts attractive through higher nutritional value when
consumption of the part services the plant, Smith (Chap. 5) and Garber and Porter
(Chap. 4) argue that pod gums should be more digestible and attractive when
compared to trunk gums because pod gums are a reward to seed dispersers.
However, we really do not yet know if these supposed differences are real for the
exudate consumer.
Power (Chap. 2) makes an important distinction between food (what is consumed)
and nutrients (what the animals need and actually get out of the food). He points out
that chemical analyses can be misleading if analytic methods are not fully understood,
that some analyses do not reflect the results of digestion, especially where fermen-
tation is involved, and that they do not tell you whether potentially available energy
is actually used. Only feeding trials can do this. However, some field observations
may be suggestive of energy payoffs of gum. Génin et al. (Chap. 6) show a lack of sex
difference in M. griseorufus gum feeding but that females are both heavier and feed
more on fruit than males. This suggests that the energy payoff in favor of females
comes from fruit, not gum. We need to be more critical “consumers” of chemical
analyses of gums as we attempt to make interpretations of their digestibility and
quality as a food resource.
Because of the digestive challenges presented by gums, differences in gut anatomy
and kinetics have been associated with differences in gum use (see below).
Heymann and Smith (1999) found that wild tamarins concentrated gum consump-
tion late in the day. This was interpreted as behavior which held the gum in the gut
during the night when defecation was less common so that it could be fermented
longer. Smith (Chap. 5) replicates and extends a previous finding from the same
site showing that tamarins consume trunk gum but not pod gum more in the late
afternoon, when it is likely to be retained in the gut overnight. Interestingly, this
would seem to be a “harvesting adaptation” that is associated with a nonpreferred,
probably fallback, food (contra Marshall and Wrangham 2007). In contrast, Porter
et al. (2009) did not find the temporal patterning of trunk and pod gum consumption
in Callimico to correspond to the predictions that pod gums would be more digestible
and thus eaten more in the morning. They found a pattern of trunk and stilt gums
being eaten more in the morning and pod gums being eaten more in the afternoon.
Génin et al. (Chap. 6) reverse the argument by suggesting that gums eaten by mouse
lemurs are readily digestible because they are consumed throughout the active
period, i.e., the night (but see below, and Powers, Chap. 2, concerning gut kinetics
of liquids vs. solids).

111 Introduction: Advances and Remaining Sticky Issues
Measuring Distribution and Seasonality of Availability
and Its Consumption
The challenges of measuring exudate consumption in ways that are comparable
across species and populations are not new to primatology (Altmann 1974;
Ray 2007). Examples of the differences in approaches are found by contrasting the
chapters in this volume by Génin et al. (Chap. 6), Garber and Porter (Chap. 4), Joly-
Radko and Zimmermann (Chap. 7), and Smith (Chap. 5) for different species and
sites. Sampling methods, definitions of behavioral categories, and specific dependent
variables used rarely are directly comparable. Some studies assess changes
(e.g., seasonal) in use of different exudates without assessing availability independently
of use. However, the latter is critical for testing various hypotheses about anatomical,
social, and cognitive correlates of food availability.
Exudates are quite challenging when it comes to measuring availability and
quality. Distribution of food patches also varies on spatial scales, which can range
from an intercontinental scale down to variation within a single plant. Distribution
can also vary on a time scale ranging from years to the rate of renewal at a single
glob. Despite these challenges several chapters do offer insights about the availability
of exudate foods and the consequences of those patterns of availability (Génin et al.,
Chap. 6 and Joly-Radko and Zimmermann, Chap. 7). Although Garber and Porter
(Chap. 4) have countered the notion that nongougers are aseasonal in use of gums,
they do suggest that tamarins and callimico may have different cognitive skills to
track the differing availability (renewal rates) and locations of their exudate
resources. Like Nekaris et al. (Chap. 8) and Génin et al. (Chap. 6), their studies
focus on renewal rate with various experimental approaches, though the methods
used varied widely across these studies as did the results and conclusions. Clearly,
some form of standardization against which to judge experiments on exudate
renewal time is needed (hourly? daily? longer periods?).
The difficulties in assessing gum availability and the factors that might influence
foraging on exudates are illustrated by the limited success in explaining inter-
population differences among pygmy marmosets in the time spent feeding on
exudates and in the specific exudate species used. Yépez et al. (2005) found that the
time spent feeding on gum did not correlate with the number of exudate species
available nor the abundance of exudate trees in each area. Within each population,
an exudate species’ relative abundance was unrelated to its relative time of con-
sumption and the number of species consumed was unrelated to group size, range
size, or amount of sampling effort. Other factors proposed which might relate to the
consumption differences were the chemical (nutritional) content of exudates,
the hardness of the wood (increasing gouging time), the viscosity of the exudates,
and degree of human disturbance of the animals.
Approaches to measuring seasonality of exudate resources and patch sizes are
quite variable. Génin et al. (Chap. 6) measured distribution of whole gum trees as
patches while Garber and Porter (Chap. 4) tried to monitor individual natural and

12 L.T. Nash and A.M. Burrows
artificial gum “sites” within trees. Isbell (1998) went further in her study of patas
monkey gum use as she measured the height of each tree, and, when gum was present,
the number of gum sites and their heights in the tree. She also visually estimated
the surface area of each gum site, and, for globular gum, the volume. Clearly, there
is much yet to be done to assess why, where and when primates consume gum.
Anatomical Adaptations to Exudativory
Just what does it mean to be a “gum specialist?” Is this defined on the basis of the
amount consumed, the seasonality of use, or anatomical specializations that relate
to it? Across chapters in this volume criteria are used in varying combinations and
sometimes leading to contradictory classifications of different species. For example,
Génin et al. (Chap. 6) seem to use year-round consumption at a high level
(diet 75% gum) as a criterion for specialist. Other chapters imply that there must
be specific anatomical features associated. In a related vein, Rosenberger (Chap. 14)
discusses the continuing debate about the role of frequency of use vs. “critical function”
use in driving the evolution of anatomical features associated with exudate acquisition
and consumption.
Cranial and Dental Anatomy: The Problem
of “Gouge” vs. “Scrape”
As a dietary niche exudativory presents two large challenges: accessing exudates and
digesting them. In 1986 we seemed to have a fair understanding of how animals
accessed exudates: callithrichines made use of their “short-tusked” anterior dentition
to gouge tree trunks and elicit exudate flow, and the exudativorous strepsirrhines
made some kind of use of their toothcomb to access exudates. However, chapters in
this book reveal an entirely less clear picture of how the strepsirrhine exudate-
feeders (which now include added taxa such as lorises and mouse lemurs) use their
dentition in acquisition (Nekaris et al., Chap. 8; Burrows and Nash, Chap. 11;
Stephenson et al., Chap. 12). Génin et al. (Chap. 6) document M. griseorufus scraping
bark to stimulate gum flow. Nekaris et al. (Chap. 8) describe in exemplary detail the
sights (and sounds) produced by Nycticebus gum-gouging behavior. They suggest
that differences in gum feeding may be associated with some of the variation in body
size and craniodental morphology within Asian lorises. They report that the tooth-
comb may be used by Nycticebus to gouge and scrape gum and that these animals
may chew on strands of gum with their molars. In contrast, among galagos, Burrows
and Nash (Chap. 11) indicate that the toothcomb may not be an important acquisition
tool in galagos, but that the more posterior dentition may be more useful.
Part of the difficulty in understanding dental characters and acquisition behavior
in strepsirrhines is that we do not have clear detailed descriptions of how these taxa

131 Introduction: Advances and Remaining Sticky Issues
use their teeth to gather gum. Nekaris et al. (Chap. 8) provide the kind of detailed
descriptions for some of the Asian lorises that are needed for other species.
They provide photographs of the holes some species gouge and descriptions of the
sounds of teeth working on branches that make it clear that gouging is certainly
occurring. Particularly strong “gouging” is sometimes described for Phaner,
Euoticus, and Allocebus, but the descriptions we have of actual acquisition behavior
and which teeth are used are minimal (see below). Among lemurs, galagos, and
Asian lorisids, it is likely that there is behavioral variability across species which
could easily be described as gouge, scoop, puncture, prise, scrape, or lick as well
as the vague “collect,” “glean,” or “acquire.” Clearly, for these nocturnal forms it
will be more difficult than with the diurnal callithrichines to get good, ideally quan-
tified, behavioral data. With available low-light video cameras, it may be possible.
Such data are critical to solving the problem of possible dental morphological
“adaptations” to exudate-feeding.
An anecdote from personal experience of LTN with captive G. senegalensis may
be useful. They were offered a paste of Acacia gum on a fingertip. First they licked
extensively and with a very long tongue extension. As the gum became depleted,
they gently scraped along the finger, but probably not hard enough to break really
hard dried gum. However, the very similar G. moholi will leave scrape marks from
its toothcomb after some feeding activities (Bearder and Martin 1980a). Rosenberger
(Chap. 14) discusses the debate about the overall function of the toothcomb, how it
may have functioned in the earliest primates that possessed one, and whether a
toothcomb itself is a reliable indicator of exudate-feeding in the fossil record.
One of the challenges to our understanding of the dental and cranial characters
involved with exudate acquisition, both in callithrichines and in strepsirrhines, is
the following question: Does gouging and/or scraping require the generation of
high forces at the anterior dentition? Yes, (Dumont 1997) and no (Vinyard et al.
2003; Taylor et al. 2009) and maybe (Burrows and Smith 2005). Dumont (1997)
found several cranial characteristics in both strepsirrhine and callithrichine exuda-
tivores consistent with generating high forces, but Vinyard et al. (2003) found few
characters associated with generating high forces. Instead, they found features con-
sistent with generating an increased gape size specifically in callithrichines. Using
the exudativorous galago O. crassicaudatus and the frugivorous Otolemur garnettii,
Burrows and Smith (2005) found a mixed bag with some characters consistent with
generating a high force at the anterior dentition and some consistent with generating
a larger gape. Recently, Taylor et al. (2009) have found fiber characteristics of the
temporalis and masseter muscles in gouging callithrichines associated with generating
a large gape size but not high forces. In the 2008 IPS symposium, Taylor, Vinyard
and White presented data comparing the trigeminal nuclei (which give rise to the
trigeminal nerve, the motor supply of the temporalis and masseter muscles) in
Saquinus oedipus and Callithrix jacchus. These results suggested that gouging may
affect the size of the proprioceptive pathways but not the motor pathways associated
with the jaws.
Several chapters in this volume bring the question of “increased force” vs.
“increased gape” to the forefront (Ravosa et al., Chap. 9; Mork et al., Chap. 10).

14 L.T. Nash and A.M. Burrows
Ravosa and colleagues present a review of the cranial characters associated with
primate exudate-feeding in both callithrichines and strepsirrhines. They synthesize
results from many studies to demonstrate that there indeed are some features in the
skulls of exudate-feeders associated with generating a greater force at the anterior
dentition and that some are associated with generating an increased gape. They also
demonstrate the anatomical trade-offs involved in generating higher forces and in
generating a larger gape but they point to a greater body of evidence supporting the
importance of the latter at the anterior dentition in exudativorous primates.
Down at the microanatomical level of the skull, Burrows and Smith (2007)
examined the cartilaginous structures of the temporomandibular joint (TMJ) in
O. crassicaudatus and O. garnettii. They found characteristics that suggested the
ability to withstand compressive forces at the TMJ articular cartilage that may be
associated with exudate-feeding. In a similar study that used gouging and nongouging
callithrichines, Mork et al. (Chap. 10) found a mix of microanatomical characters
in the TMJ articular cartilage that are both consistent and inconsistent with expectations
of how the articular cartilage would be loaded at large gapes. Their findings along
with those of Burrows and Smith (2007) reinforce the mosaic nature of morpho-
logical associations with exudativory and the potential pitfalls of inferring behavior
based upon morphological characters. In the current volume, both Ravosa et al.
(Chap. 9) and Mork et al. (Chap. 10) point to the need for more species to be studied
for both behavior and morphology and the value of ontological studies of morphology.
Rosenberger (Chap. 14) presents a hypothetical model of morphological characters
that might be related to the biomechanical problems associated with gouging and
scraping (“gleaning” in his terminology) exudates and how these characters may be
reflected in the primate fossil record. He uses plesiadapiforms in his discussions of
the earliest primate and evaluates the potential role exudate-feeding may have had
in primate evolution and in the lifestyles of the plesiadapiforms. In his chapter,
Rosenberger makes a plea for the use of tooth wear as a characteristic of considerable
weight when evaluating morphological signals of exudativory. Rosenberger’s chapter
continues, and certainly does not solve, the debate about the functional reasons
toothcombs evolved (diet, and which diet, or social, i.e., grooming, behavior).
One of the key pieces of data missing from a more complete answer to this question
is material properties and hardness of the plants being gouged and/or scraped. As was
found with different parts of bamboo, which sometimes have different mechanical
properties and sometimes did not (Yamashita et al. 2009), different kinds and locations
of gum may produce different mechanical effects that need to be accounted for in
comparative work on morphological adaptations associated with exudate-feeders.
As is apparent, one of the problems we face is a proper accounting of the acqui-
sition behavior. A problem in the literature is that behaviors hypothesized based on
anatomy in an original citation became “reified” in a secondary source which states
the behavior happens and then a third source cites the second. This becomes analo-
gous to the children’s game Americans call “playing telephone” or “gossip”
(see http://www.Wikipedia.org “telephone game”) where a message is more garbled
the more people it passes through. For example, in describing Phaner, Euoticus,
and Allocebus, LTN stated.

151 Introduction: Advances and Remaining Sticky Issues
Phaner and Euoticus, the most gummivorous lemuroid and lorisoid, respectively, possess
a caniniform first upper premolar. This has been suggested as an adaptation for gouging,
but the detailed behavioral observations needed to confirm this function are lacking
(Charles-Dominique 1977; Charles-Dominique and Petter 1980). A. trichotis, a cheir -
ogalid, also shows this dental trait. It is known only from two museum specimens (Tattersall
1982). Based on head lengths of the specimens, it is about the size of G. senegalensis or
Cebuella. Unfortunately, this species is exceedingly rare or extinct (Tattersall 1982), so that
the extent of gummivory in its dietary adaptation may not be established with certainty.
Nash (1986, p. 125)
This was based on the following, which are all that are provided by Charles-
Dominiques’ direct observations of Euoticus or Phaner: “The author has…
(observed) Euoticus… visiting certain wound areas on trees where tiny droplets of
gum were forming, and collecting the exudate by licking or scooping with the
toothscraper.” (Charles-Dominique 1977, p. 42) and
Phaner… uses its long tongue and also the narrow and procumbent tooth comb to scoop the
gum which would otherwise be inaccessible without such a “tool.” Phaner furcifer is also
characterized by the development of the upper first premolar (caniniform). This peculiarity,
seen only in A. trichotis, among Malagasy lemurs and Galago ( Euoticus) elegantulus, among
African Lorisids, appears to be an adaptation for the extraction of vegetal exudations.
Charles-Dominique and Petter (1980, p. 78)
Nevertheless, some authors use Nash (1986) as the citation to support that Phaner,
Euoticus or even Allocebus gouge into bark and are known to have a diet high in
gouged gums – “On the one hand, they [gums] account for the major part in the diet
of P. furcifer (65%) and A. trichotis.” (Viguier 2004, p. 496). Others recognize that
dietary interpretations of Allocebus are based on morphological analogy to Euoticus
(e.g., Masters and Brothers 2002). We now have the very recent field work of
Biebouw (2009), which indicates that a major part of the diet of Allocebus is,
indeed, gum. The details of how they acquire it, though, are yet to be described. The
same is true for Phaner and Euoticus.
Soft Tissues: Guts, Tongues, and Pelage
The chemical structure of exudates, and certainly gums, presents digestive chal-
lenges (Power, Chap. 2). If they must be fermented in the digestive tract, we also
expect, and have found, that the size, proportions, and kinetics of the gut are associ-
ated with the degree of gummivory in primates. Gum needs to remain in the gut for
a longer time vs. other foods in order to be fermented. Power points out that the
relationships among gut morphological variables and gut kinetics are complex.
Lambert (1998) noted that there is only a weak relationship between gut transit time
and body size. Gut transit time is influenced by the competing problems of absorp-
tion vs. processing (Power, Chap. 2). Another problem is to what extent primates
show “modularity,” i.e., the ability of an individual or species to regulate digestion
relative to the current diet (Lambert 1998).

16 L.T. Nash and A.M. Burrows
The best data we have on digestive adaptations of the gut, as with other
morphological features, comes from contrasts within the callithrichines (Power,
Chap. 2). Marmosets seem to have more capacious hindguts than tamarins. Power
emphasizes the differences in how the anatomy and activity of the gut can vary
independently of each other. Measuring gut kinetics can be tricky, as it is ideal to
have separate markers for the solid and fluid fractions of the digesta. In addition,
different size solid markers may give different results, complicating comparative
studies. This is important because it has been hypothesized that gum eaters may
have a “cecal-colonic separation mechanism” that selectively retains the gum
(which is soluble) in the gut relative to solids (e.g., seeds, insect exoskeletons)
(Caton et al. 1996, 2000). Captive G. moholi that are fed a diet mostly of gum show
the expected slower gut transit time of the fluid digesta compared to the solid fraction.
The ecologically similar G. senegalensis has been shown to slow overall transit rate
(as indicated by solid markers) when shifted from a diet high in fruit to one high in
gum (Nash 1989).
Power indicates the complexity of unraveling the adaptations when he shows
that within callithrichines there may not be simple correlations between proportion
of gum eaten and gut kinetics. His work on marmosets and tamarins demonstrates
different gut adaptations within gum consumers depending on the other foods
(insects vs. fruits) consumed. Some combinations may present a diet within which
there are items that would be optimally handled by the gut in different ways, pre-
senting conflicting adaptive challenges to be solved. This recalls the issue of trade-offs
in the craniofacial anatomy that may be associated with exudate use (see above).
However, until we have comparable data for a wider variety of exudativores on gut
anatomy, gut kinetics on different diets, and studies of net energy gain from feeding
studies, we will not have a full understanding of primate digestive adaptations to
the challenges of exudates.
Another two areas that are still in need of more systematic study are tongue
anatomy and features of the pelage. Génin et al. (Chap. 6) and Nekaris et al.
(Chap. 8) both present data that associate relatively long tongues with exudativory
(see also LTN’s anecdote, above). Nekaris et al. echo Garber (Garber 1980) in
suggesting that patterns of dorsal strips on lorises (some of which appear only
seasonally) and color patterns in some callithrichines might help camouflage animals
in exposed locations as they concentrate on gum acquisition. These notions beg for
experimental studies of the hunting behavior of these primates’ predators using
classical ethological approaches. In addition, examination of pelage features as
camouflage in other trunk forages, e.g., colugos, vs. similar-sized nontrunk forages
would help test the notion that trunk foragers have evolved pelage that hides then
while they are particularly vulnerable.
While long tongues may help in acquiring gums, exudate use may also influence
taste sensitivities though there is little information on taste thresholds to specifically
look at the effects of exudates (Nash 1989; Simmen and Hladik 1998). Such studies
are complicated by allometric effects and the patterning of requirements to avoid
secondary compounds. Docherty et al. (Chap. 13) address detailed features of
tongue papillae, the area of the tongue holding taste receptors, in Otolemur that
may relate to taste sensitivity. These authors suggest that the frugivorous O. garnettii

171 Introduction: Advances and Remaining Sticky Issues
has a greater density of taste receptors than the exudativorous O. crassicaudatus
and that this difference is consistent throughout ontogeny. Results of this study may
speak to the chemical signals associated not only with finding exudates but life-
history variables linked to exudate-feeding.
Locomotion and Limbs
At the 2008 IPS symposium Ford presented findings that indicated there were few
if any shared postcranial features across all gummivorous primates even though
others have implicated a link to keeled, pointed, or claw-like nails. She noted the
need for better studies of positional behavior in such species and their nongum-
mivorous sister taxa that incorporated feeding and other contexts and for more
biomechanical work on the problem. A practical problem for such work is the rarity
of skeletons of some important taxa in museum collections (e.g., Phaner, Cebuella,
Mico) and of captive colonies of living animals. Like the Taylor et al. work on the
trigeminal nuclei, we look forward to the publication of this work elsewhere.
Nekaris et al. (Chap. 8) present excellent qualitative descriptions of some of the
positional behaviors associated with exudate eating in different lorises. She notes
they use head-down postures extensively though they do not have keeled nails. This
calls attention to the need to distinguish movement on angled substrates that is
head-up vs. head-down (Crompton 1983; Harcourt and Nash 1986b) as this may
influence the grip problems associated with gum foraging. Génin et al. (Chap. 6)
note that most cheirogalieds have pointed nails. Garber and Porter (Chap. 4) point
out differences between callimicos and tamarins in positional behavior during for-
aging on pod vs. trunk exudates. Both use vertical clinging and leaping to get to
trunk gums but callimicos only use pods that have fallen to the ground while tama-
rins can hang by their feet in the canopy to harvest pods.
Stephenson et al. (Chap. 12) examine volar pad and nail features among galagos.
They confirm previous studies (Anderson 1999; Anderson et al. 2000) that indicate
volar pad size and shape may be related to taxonomic issues and body size.
Unfortunately, no clear association with diet is clear in their analyses for either pads
or nail shapes. This may again be related to the comparative samples chosen and
the need for more detailed information on the substrates and positional behaviors
used in food acquisition. It may not be the food per se, but the postures and sub-
strates that are important, as has been suggested for the evolution of claw-like nails
in callithrichines (Garber et al. 1996).
Diet and Captive Husbandry of Exudativores
Earlier data on primate exudativory have stimulated attempts to use natural
branches and artificial gum feeders as enrichment devices for captive animals,
mainly marmosets (Kelly 1993). This work has also demonstrated management

18 L.T. Nash and A.M. Burrows
benefits; as animals come close to keepers offering gum in feeders, they can be
examined for health and welfare. In addition, some feeders provide stimulation and
favorably influence activity patterns. Recently, Huber (2009a, b) surveyed a variety
of zoos worldwide to see how gum was used as enrichment and/or a dietary
supplement. The majority of responding zoos housing marmosets provided them
with gum, but far fewer did for exudativorous tamarins and galagos. Apparently the
prominence of gouging behavior influenced management decisions. Huber (2 009a, b)
also found that zoos do not appreciate field studies showing that the diet of patas is
high in gum. She also reviews methods of presenting gum that emulate natural
foraging problems and stimulate more naturalistic positional behavior. As always,
there are trade-offs in management decisions. Power (Chap. 2) worries that if gum
is fed as enrichment but captive animals then eat less of their carefully balanced diet
of other “normal” foods, extra calcium might be ingested at the expense of vitamins
from fruits. He suggests that there is no “demonstrated nutritional reasons” for giving
captive primates gum beyond behavioral enrichment. In contrast, Nekaris et al.,
(Chap. 8) point out that providing gum-gouging opportunities may be important in
captive management of lorises, as it may improve dental health and increase activity
to limit obesity, both of which are common problems of captive lorises.
Concluding Remarks
It now seems clear that exudativory has evolved multiple times within primates,
although the issue of whether it was a feature in the earliest primates remains unre-
solved, and possibly unresolvable (Rosenberger, Chap. 14). Compared to 1986,
it seems less clear that exudativory was ancestral within all callithrichines. The vari-
ability within both marmosets and tamarins suggests that the morphological differences
between them would be profitably examined at a finer scale. As we learn more
about exudativory, the diversity of both morphological and behavioral features that
are associated with it in different types of primates are likely to become more complex.
The ability to generalize from the better known callithrichine features to other
primates mostly remains to be established. It is not entirely clear that the cranio-
dental morphological differences between the gouging and nongouging callithrichines
will be informative about the contrasts among strepsirrhines that vary in their use
of gums. Similarly, the evolution of the digits (claws, keeled or pointed nails) does
not easily “track” patterns of exudativory.
The work in this volume highlights several directions for future work. It is clear
that more detailed observations are needed in the field of how strepsirrhines actually
use their mouth and dentition to acquire gum. Unfortunately, a major limitation on
work in strepsirrhines will be their scarcity in captivity where biomechanical studies
are more feasible. Also, more work on the costs and benefits of exudates as enrichment
for captive primates is needed.
We also need more detailed fieldwork on all the exudativores to allow comparative
studies of how the distribution of their foods in time and space influences social and

191 Introduction: Advances and Remaining Sticky Issues
cognitive evolution. Here we include not only primates, but the few other mammals
that eat exudates such as sugar gliders (see above) and some didelphids (Génin et al.,
Chap. 6). Such work is predicated on knowing better how exudates are distributed
and what they really provide in the way of nutrition. A more complete understanding
of the temporal and spatial availability of this food resource is required to under-
stand how it is adapted to behaviorally. Are there continental differences in the role
of primate exudativores in their communities (Reed and Bidner 2004)? More
detailed understanding of the role of exudates in primate diets can more fully
inform conservation efforts for these species, too many of which are rapidly disap-
pearing. If exudativory is indeed an adaptation to hypervariable environments, will
primate exudativores fare better in the face of climate change (Wright 1999) or will
the effects of exudativory on species survival also be “clade dependent?”
Acknowledgments First, we thank all of the authors whose hard work tackling the sticky issue
of exudativory – and their patience with our nagging – make this an exciting volume. To all the
reviewers of the chapters in the book, we are most grateful for the improvements they helped us
make to the chapters. We are very grateful to Melissa Higgs at Springer for her endless advice and
assistance. Michael Power and George Perry provided helpful comments on this chapter. We
would also like to thank our families who put up with our rants and occasional absences as well
as providing us with much needed support.
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25
Abstract Gum is an unusual food that presents significant challenges to animals
that feed on it. Gum is limited in availability; trees generally secrete it only in
response to damage. Gum is a b-linked complex polysaccharide, and as such is resistant
to mammalian digestive enzymes and requires fermentation by gut microbes. It
contains little or no lipid, low amounts of protein, and no appreciable levels of vita-
mins. As a food, gum can be characterized as difficult to obtain, potentially limited
in quantity, difficult to digest, and primarily a source of energy and minerals. Despite
these drawbacks, many primates feed extensively on gums. Among mammals,
gum-feeding largely appears to be a primate dietary adaptation. Why are there so
many primate gum-feeders and what adaptations have allowed them to make a
living on such a problematic food? This is the central question of this book. This
chapter examines digestive and nutritional aspects of gum. Specific examples of
biological adaptations found in common and pygmy marmosets (Callithrix jacchus
and Cebuella pygmaea), small New World primate gum-feeding specialists, will
be examined. These marmoset species have many similarities in their behavior,
morphology and metabolism, but also some instructive differences in their diges-
tive function. C. pygmaea is the smallest of the marmosets but has the slowest
passage rate of digesta. This might represent an adaptation to retain difficult-to-digest
material, such as gum, within the gut to allow fermentation. In contrast, C. jacchus
has a rapid passage rate. Passage rate in C. jacchus appears adapted more for rapidly
excreting indigestible material (e.g., seeds) than for retaining gum within the gut
to enable more complete digestion. Fruit is a rare component of C. pygmaea’s
diet; hence any constraint on feeding caused by filling the gut with ingested seeds
is greatly relaxed, apparently enabling digestive kinetics that favor digestive effi-
ciency over maximizing food intake. Interestingly, however, these marmosets share
M.L. Power (*)
Nutrition Laboratory, Smithsonian Conservation Biology Institute, National Zoological Park,
P.O. Box 37012, MRC 5503, Washington, DC 20013-7012, USA
and
Research Department, American College of Obstetricians and Gynecologists,
Washington, DC 20024, USA
e-mail: [email protected]
Chapter 2
Nutritional and Digestive Challenges to Being
a Gum-Feeding Primate
Michael L. Power
A.M. Burrows and L.T. Nash (eds.), The Evolution of Exudativory in Primates,
Developments in Primatology: Progress and Prospects,
DOI 10.1007/978-1-4419-6661-2_2, © Springer Science+Business Media, LLC 2010

26 M.L. Power
an ability to digest gum despite their differences in gum kinetics. In captivity both
species have been shown to be more able to digest Acacia gum than related species
that feed less often on gum in the wild.
Introduction
All life has a common biochemical underpinning. Because of this, everything living
potentially is food. Indeed, for every living organism there are other organisms that
feed off of it. However, because of the immense amount of time over which life has
diverged and radiated, the common biochemical underpinning has accumulated a
tremendous amount of variation in specific characteristics among taxa. Everything
may be food for something; but for any given organism most of what is in its
environment is not food.
Animals eat food; they require nutrients. A significant proportion of anatomy
and physiology has as its primary purpose the transformation of food that animals
select from their environment into the nutrients required for life. These challenges
can be external ones such as finding and acquiring food, and they can be internal
challenges, such as digesting, assimilating, and metabolizing food, and then finally
excreting the associated waste products (Chivers et al. 1984). My research focus is
on the internal challenges different foods provide.
All foods provide challenges; there is no perfect food. Different foods provide
different challenges. For example, carnivores are confronted with very different
challenges in obtaining nutrients than are herbivores. Animals and plants share an
evolutionary history, and thus are biochemically similar; however, they are also
very different, reflecting the billions of years of evolutionary separation. Thus, in
general it is assumed that carnivores are faced with a less difficult nutritional challenge
than are herbivores. If you are what you eat, then eating other animals should provide
fewer difficulties than eating plants.
As is true for most generalities in biology, the one above is an oversimplification.
Although other animals certainly contain all the nutrients an animal needs to consume,
they do not contain them in the correct proportions. Animals contain far more protein
than is necessary for another animal to consume, and far less glucose and other
carbohydrate than is needed to survive. Strict carnivores must deal metabolically
with an excess of protein and insufficient carbohydrate. For herbivores the situation
is possibly reversed. Protein can be a limiting nutrient but carbohydrate is usually
in plentiful supply, though not always in a readily metabolizable form. Many plant
carbohydrates are difficult to digest. Animals that feed largely on plant material
generally face greater digestive challenges; for strict carnivores the challenges are
primarily metabolic.
Of course all plant foods are not alike, and thus provide different digestive and
metabolic challenges. This essay concerns the challenges presented by a rather
unusual plant food, gum, a type of exudate produced by certain trees and lianas.
The number of animals known to regularly feed on tree exudates is not large,

272 Nutritional and Digestive Challenges to Being a Gum-Feeding Primate
though the list is slowly expanding. Among mammals, the primate order contains
a considerable number of species that utilize tree exudates as a food resource,
including many that appear to specialize on exudates. The only other (known)
exudate-feeding mammals are a guild of small marsupials native to Australia
(Hume 1982; Smith and Lee 1984), where there are no native nonhuman primates.
Why are there so many primate gum-feeders and what adaptations have allowed
them to make a living on such a problematic food?
That is the interesting question that has inspired this book. My contribution will
be limited to examining the nutritional, digestive and metabolic advantages and
challenges from eating gum, and is further constrained by a focus on the biology of
the marmosets, small gum-feeding New World primates. Hopefully this chapter
will provide a broad enough context that the value of gum as food for other species
can be evaluated as well.
This chapter will start with a general assessment of gum as food, with some
comparison to other plant foods including other exudates. Gum is dietary fiber; as
such it presents digestive difficulties. The advantages and disadvantages of fore-
and hind-gut fermentation for obtaining nutrients from gum are briefly reviewed.
The chapter then focuses on marmosets, specifically their digestive function and
how that may or may not be adapted to gum-feeding.
Exudates as Food
There are primate species, our own especially, that incorporate a substantial amount
of animal matter in the diet; but in general, primates feed predominantly on plant
foods. Plants are complex structures. A tree is composed of many different parts
that vary widely in chemical and physical composition. Its wood, bark, leaves,
flowers, fruits, and exudates all provide food for something but any given animal
species generally will feed only on certain parts and will ignore the rest. A tree may
have many species visiting it, each feeding on a different tree product. These are
obvious statements; the point is that different plant products need to be categorized
in ways that reflect the kind of nutrition they provide in order to explore the dietary
adaptations of our subject species. To say that gum is a plant product doesn’t help
to determine what nutrition it can or cannot provide for a species. Ideally gum
should be chemically assayed to determine its constituents, and then fed to animals
in controlled trials to determine the bioavailability of those constituents. However,
there are general principles that can be used to predict what nutritional category a
plant product is likely to occupy.
There are many ways to categorize plant foods. For the purposes of this essay
I propose two simple categorizations: alive vs. not alive, and primarily reproductive
vs. primarily nonreproductive. The first categorization separates exudates from
other plant foods such as leaves, flowers, and fruit in a fundamental way. Exudates
are created by living things but they themselves are not alive. More to the point,
exudates do not contain living cells. Cells, by necessity, contain the required chemical

28 M.L. Power
components for life. Foods such as leaves, flowers and fruit that are composed
predominantly of living cells in theory should provide excellent nutrition. Of course,
there might be some difficulty in accessing the cell contents and obtaining the
nutrients; but they are there. Exudates are created by living things and therefore it
is not surprising that they contain nutrients; however, there is no expectation that
exudates will contain all the nutrients necessary for life. And indeed they do not.
Gums and other exudates are not complete foods.
Plants produce a number of exudates; in this essay I will be primarily concerned
with gum; however, the other types of exudates are briefly considered here. Tree
exudates are generally categorized as sap, gum, latex, and resin (Nash 1986). In
addition, nectar can be considered a plant exudate.
All these exudates have different functions, which influence their characteristics
as food. Consider nectar, an exudate produced by flowering plants. Flowering
plants arose more than a 100 million years ago (Soltis and Soltis 2004), greatly
diversified in the Cretaceous, and have been coevolving with animals ever since.
Nectar serves as a reward to pollinators. The plant provides food in exchange for
assistance in sexual reproduction. Nectar has evolved to be food. Therefore, it is not
surprising that nectar has characteristics that make it edible and that it provides
some nutrition, in most cases primarily energy.
These are characteristics of many, but not all plant parts that primarily serve a
reproductive function. Nectar and fruit are the principal examples. In both cases the
plant produces a food-like substance to reward animals that assist in the plant’s
reproduction. But seeds also can provide high quality nutrition. It is true that for
many plants it is beneficial if their seeds are ingested; but not if the seeds are actually
digested. There are seed dispersers and there are seed predators. For the predators,
seeds are food and quite good food for the basic reason that seeds must contain
most if not all the molecules necessary for life. They are incipient life.
Most exudates other than nectar generally don’t serve the reproductive needs
of plants, and have not evolved to be food. Sap perhaps is closest to nectar in con-
stitution. Sap contains the simple sugars from photosynthesis, and other nutrients
that are required by the plant cells to survive. The main difference between sap
and nectar is that nectar is concentrated. There are animals (mainly insects) that
feed on sap. The challenges sap presents are mainly related to acquiring it in the
first place; however, it is dilute and thus a large quantity of water must be
ingested to provide a fairly small amount of nutrition. Cicadas are an insect that
feeds on sap; the author has walked through the mist created by millions of
cicadas feeding on tree sap, and necessarily excreting large amounts of water
into the air. Joly-Radko and Zimmermann (Chap. 7) describe “sap eating by
proxy” in mouse lemurs consuming the excretions produced by hemipteran
insects feeding on sap.
Resins are phenol and terpene derivatives. They are generally considered noxious
and even toxic. There are animals that are tolerant of resins, however. The desert
wood rat (Neotoma lepida) feeds extensively on creosote bush leaves, at least in
certain areas of its range (Mangione et al. 2000). These leaves can contain as much
as 25% of the dry mass as phenolic resin (Rhoades and Cates 1976).

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conducted a campaign worthy of Henry Clay. Multitudes crowded to
hear and see a man candid enough to deliver his honest opinions
with the boldness of “Old Hickory.” The masses knew of Mr. Sibley’s
courage, sagacity and success in business, but they were unprepared
to find so sturdy a defender of their rights. His manly independence,
ringing denunciations of wrong, grand simplicity and incisive logic
aroused unbounded enthusiasm. The tide in favor of the fearless
advocate of fair-play for the lowliest creature no earthly power could
stem. His opponent was buried out of sight and Sibley was elected
by a sweeping majority.
Mr. Sibley’s course in Congress amply met the expectations of his
most ardent supporters. The prestige of his great victory, added to
his personal magnetism and rare geniality, at the very outset gave
him a measure of influence few members ever attain. During the
extra-session he expressed his views with characteristic vigor. A
natural leader, close student and keen observer, he did not wait for
somebody to give him the cue before putting his ideas on record. In
the silver-discussion he bore a prominent part, opposing resolutely
the repeal of the Sherman act. His wonderful speech “set the ball
rolling” for those who declined to follow the administration program.
The House was electrified by Sibley’s effort. Throughout his speech
of three hours he was honored with the largest Congressional
audience of the decade. Aisles, halls, galleries and corridors were
densely packed. Senators came from the other end of the Capitol to
listen to the brave Pennsylvanian who dared plead for the white
metal. For many years Mr. Sibley has been a close student of political
and social economics and he so grouped his facts as to command
the undivided attention and the highest respect of those who
honestly differed from him in his conclusions. Satire, pathos, bright
wit and pungent repartee awoke in his hearers the strongest
emotions, entrancing the bimetalists and giving their enemies a cold
chill, as the stream of eloquence flowed from lips “untrained to
flatter, to dissemble or to play the hypocrite.” Thenceforth the
position of the representative of the Twenty-sixth district was
assured, despite the assaults of hireling journals and discomfited
worshippers of the golden calf.

He took advanced ground on the Chinese question, delivering a
speech replete with patriotism and common-sense. An American by
birth, habit and education, he prefers his own country to any other
under the blue vault of heaven. The American workman he would
protect from pauper immigration and refuse to put on the European
or Asiatic level. He stands up for American skill, American ingenuity,
American labor and American wages. Tariff for revenue he approves
of, not a tariff to diminish revenue or to enrich one class at the
expense of all. The tiller of the soil, the mechanic, the coal-miner,
the coke-burner and the day-laborer have found him an outspoken
champion of their cause. Small wonder is it that good men and
women of all creeds and parties have abiding faith in Joseph C.
Sibley and would fain bestow on him the highest office in the
nation’s gift.
Human nature is a queer medley and sometimes manifests streaks
of envy and meanness in queer ways. Mr. Sibley’s motives have been
impugned, his efforts belittled, his methods assailed and his neckties
criticised by men who could not understand his lofty character and
purposes. The generous ex-Congressman must plead guilty to the
charge of wearing clothes that fit him, of smoking decent cigars, of
driving fine horses and of living comfortably. Of course it would be
cheaper to buy hand-me-down misfits, to indulge in loud-smelling
tobies, to walk or ride muleback, to curry his own horses and let his
wife do the washing instead of hiring competent helpers. But he
goes right ahead increasing his business, improving his farms,
developing American trotters and furnishing work at the highest
wages to willing hands in his factories, at his oil-wells, on his lands,
in his barns and his hospitable home. He dispenses large sums in
charity. His benevolence and enterprise reach far beyond
Pennsylvania. He does not hoard up money to loan it at exorbitant
rates. As a matter of fact, from the hundreds of men he has helped
pecuniarily he never accepted one penny of interest. He has been
mayor of Franklin, president of the Pennsylvania State-Dairymen’s
Association, director of the American Jersey-Cattle Club and member
of the State Board of Agriculture. He is a brilliant talker, a profound

thinker, a capital story-teller and a loyal friend. “May he live long and
prosper!”
Miller & Sibley’s Prospect-Hill Stock-Farm is one of the largest, best
equipped and most favorably known in the world. Different farms
comprising the establishment include a thousand acres of land
adjacent to Franklin and a farm, with stabling for two-hundred
horses and the finest kite-track in the United States, at Meadville. On
one of these farms is the first silo built west of the Allegheny
mountains. Trotting stock, Jersey cattle, Shetland ponies and Angora
goats of the highest grades are bred. For Michael Angelo, when a
calf six weeks old, twelve-thousand-five-hundred dollars in cash were
paid A. B. Darling, proprietor of the Fifth-Avenue Hotel, New York
City. Animals of the best strain were purchased, regardless of cost.
In 1886 Mr. Sibley bought from Senator Leland Stanford, of
California, for ten-thousand dollars, the four-year-old trotting-stallion
St. Bel. Seventy-five thousand were offered for him a few weeks
before the famous sire of numerous prize-winners died. Cows that
have broken all records for milk and butter, and horses that have
won the biggest purses on the leading race-tracks of the country are
the results of the liberal policy pursued at Prospect-Hill. Charles
Marvin, the prince of horsemen, superintends the trotting
department and E. H. Sibley is manager of all the Miller & Sibley
interests. Hundreds of the choicest animals are raised every year.
Prospect-Hill Farm is one of the sights of Franklin and the enterprise
represents an investment not far short of one-million dollars.
Wouldn’t men like Charles Miller and Joseph C. Sibley sweep away
the cobwebs, give business an impetus and infuse new life and new
ideas into any community?
Franklin had tallied one for heavy-oil, but its resources were not
exhausted. On October seventeenth, 1859, Colonel James P. Hoover,
C. M. Hoover and Vance Stewart began to drill on the Robert-
Brandon—now the Hoover—farm of three-hundred acres, in
Sandycreek township, on the west bank of the Allegheny river, three
miles south of Franklin. They found oil on December twenty-first, the
well yielding one-hundred barrels a day! This pretty Christmas gift
was another surprise. Owing to its distance from “springs” and the

two wells—Drake and Evans—already producing, the stay-in-the-rut
element felt confident that the Hoover Well would not “amount to a
hill of beans.” It was “piling Ossa on Pelion” for the well to produce,
from the second sand, oil with properties adapted to illumination and
lubrication. The Drake was for light, the Evans for grease and the
Hoover combined the two in part. Where and when was this
variegated dissimilarity to cease? Perhaps its latest phase is to come
shortly. Henry F. James is beginning a well south-west of town, on
the N. B. Myers tract, between a sweet and a sour spring. Savans,
scientists, beer-drinkers, tee-totalers and oil-operators are on the
ragged edge of suspense, some hoping, some fearing, some praying
that James may tap a perennial fount of creamy ’alf-and-’alf.
Once at a drilling-well on the “Point” the tools dropped suddenly.
The driller relieved the tension on his rope and let the tools down
slowly. They descended six or eight feet! The bare thought of a
crevice of such dimensions paralyzed the knight of the temper-screw,
all the more that the hole was not to the first sand. What a lake of
oil must underlie that derrick! He drew up the tools. They were
dripping amber fluid, which had a flavor quite unlike petroleum. Did
his nose deceive him? It was the aroma of beer! A lick of the stuff
confirmed the nasal diagnosis—it had the taste of beer! The alarm
was sounded and the sand-pump run down. It came up brimming
over with beer! Ten times the trip was repeated with the same
result. Think of an ocean of the delicious, foamy, appetizing German
beverage! Word was sent to the owners of the well, who ordered the
tubing to be put in. They tried to figure how many breweries the
production of their well would retire. Pumping was about to begin, in
presence of a party of impatient, thirsty spectators, when an excited
Teuton, blowing and puffing, was seen approaching at a breakneck
pace. Evidently he had something on his mind. “Gott in Himmel!” he
shrieked, “you vas proke mit Grossman’s vault!” The mystery was
quickly explained. Philip Grossman, the brewer, had cut a tunnel a
hundred feet into the hill-side to store his liquid-stock in a cool place.
The well chanced to be squarely over this tunnel, the roof of which
the tools pierced and stove in the head of a tun of beer! Workmen
who came for a load were astonished to discover one end of a string

of tubing dangling in the tun. It dawned upon them that the drillers
three-hundred feet above must have imagined they struck a crevice
and a messenger speeded to the well. The saddened crowd slinked
off, muttering words that would not look nice in print. The tubing
was withdrawn, the hogshead was shoved aside, the tools were
again swung and two weeks later the well was pumping thirty
barrels a day of unmistakable heavy-oil.
The Hoover strike fed the flame the Evans Well had kindled. Lands
in the neighborhood were in demand on any terms the owners might
impose. From Franklin to the new well, on both sides of the
Allegheny, was the favorite choice, on a theory that a pool connected
the deposits. Leases were snapped up at one-half royalty and a
cash-bonus. Additional wells on the Hoover rivaled No. 1, which
produced gamely for four years. The tools were stuck in cleaning it
out and a new well beside it started at sixty barrels. The “Big-Emma
Vein” was really an artery to which for years “whoa, Emma!” did not
apply. Bissell & Co. and the Cameron Petroleum-Company secured
control of the property, on which fifteen wells were producing two-
hundred barrels ten years from the advent of the Hoover & Vance.
Harry Smith, a city-father, is operating on the tract and drilling
paying wells at reasonable intervals. Colonel James P. Hoover died on
February fourth, 1871, aged sixty-nine. Born in Centre county, he
settled in the southern part of Clarion, was appointed by Governor
Porter in 1839 Prothonotary of Venango county and removed to
Franklin. The people elected him to the same office for three years
and State-Senator in 1844. The Canal-Commissioners in 1851
appointed him collector of the tolls at Hollidaysburg, Blair county, for
five years. He filled these positions efficiently, strict adherence to
principle and a high sense of duty marking his whole career. The
esteem and confidence he enjoyed all through his useful life were
attested by universal regret at his death and the largest funeral ever
witnessed in Franklin. His estimable widow survived Colonel Hoover
twenty years, dying at the residence of her son-in-law, Arnold
Plumer, in Minnesota. Their son, C. M. Hoover, ex-sheriff of the
county, has been interested in the street railway. Vance Stewart, who
owned a farm near the lower river-bridge, removed to Greenville and

preceded his wife and several children, one of them Rev. Orlando V.
Stewart, to the tomb. Another son, James Stewart, was a prominent
member of the Erie bar.
B. E. SWAN.
The opening months of 1860 were decidedly lively on the Cochran
Farm, in Cranberry township, opposite the Hoover. The first well, the
Keystone, on the flats above where the station now stands, was a
second-sander of the hundred-barrel class. The first oil sold for
fourteen dollars a barrel, at which rate land-owners and operators
were not in danger of bankruptcy or the poor-house. Fourteen-
hundred dollars a day from a three-inch hole would have seemed too
preposterous for Munchausen before the Pennsylvania oil-regions
demonstrated that “truth is stranger than fiction.” The Monitor,
Raymond, Williams, McCutcheon and other wells kept the production
at a satisfactory figure. Dale & Morrow, Horton & Son, Hoover & Co.,
George R. Hobby, Cornelius Fulkerson and George S. McCartney
were early operators. B. E. Swan located on the farm in May of 1865
and drilled numerous fair wells. He has operated there for thirty-two
years, sticking to the second-sand territory with a tenacity equal to
the “perseverance of the saints.” When thousands of producers,
imitating the dog that let go the bone to grasp the shadow in the
water, quit their enduring small wells to take their chance of larger
ones in costlier fields, he did not lose his head and add another to
the financial wrecks that strewed the greasian shore. Appreciating

ALEXANDER COCHRAN.
his moral stamina, his steadfastness and ability, Mr. Swan’s friends
insist that he shall serve the public in some important office. Walter
Pennell—his father made the first car-wheels—and W. P. Smith drilled
several snug wells on the uplands, Sweet & Shaffer following with six
or eight. Eighteen wells are producing on the tract, which contains
one-hundred-and-forty acres and has had only two dry-holes in its
thirty-six years of active developments.
Alexander Cochran, for forty years
owner of the well-known farm bearing his
name, is one of the oldest citizens of
Franklin. Winning his way in the world by
sheer force of character, scrupulous
integrity and a fixed determination to
succeed, he is in the highest and best
sense a self-made man. Working hard in
boyhood to secure an education, he
taught school, clerked in general stores,
studied law and was twice elected
Prothonotary without asking one voter for
his support. In these days of button-
holing, log-rolling, wire-pulling, buying and soliciting votes this is a
record to recall with pride. Marrying Miss Mary Bole—her father
removed from Lewistown to Franklin seventy-five years ago—he built
the home at “Cochran Spring” that is one of the land-marks of the
town and established a large dry-goods store. As his means
permitted he bought city-lots, put up dwelling-houses and about
1852 paid sixteen-hundred dollars for the farm in Cranberry township
for which in 1863, after it had yielded a fortune, he refused seven-
hundred-thousand! The farm was in two blocks. A neighbor
expostulated with him for buying the second piece, saying it was
“foolish to waste money that way.” In 1861, when the same neighbor
wished to mortgage his land for a loan, he naively remarked: “Well,
Aleck, I guess I was the fool, not you, in 1852.” A man of broad
views, Mr. Cochran freely grants to others the liberality of thought he
claims for himself. A hater of cant and sham and hollow pretence, he
believes less in musty creeds than kindly deeds, more in giving

loaves than tracts to the hungry, and takes no stock in religion that
thinks only of dodging punishment in the next world and fails to help
humanity in this. In the dark days of low-priced oil and depressed
trade, he would accept neither interest from his debtors nor royalty
from the operators who had little wells on his farm. He never
hounded the sheriff on a hapless borrower, foreclosed a mortgage to
grab a coveted property or seized the chattels of a struggling victim
to satisfy a shirt-tail note. There is no shred of the Pecksniff, the
Shylock or the Uriah-Heep in his anatomy. At fourscore he is hale
and hearty, rides on horseback, cultivates his garden, attends to
business, likes a good play and keeps up with the literature of the
day. The productive oil-farm is now owned by his daughters, Mrs. J.
J. McLaurin, of Harrisburg, and Mrs. George R. Sheasley, of Franklin.
The proudest eulogy he could desire is Alexander Cochran’s just
desert: “The Poor Man’s Friend.”
Down to Sandy Creek many wells were drilled from 1860 to 1865,
producing fairly at an average depth of four-hundred-and-fifty to
five-hundred feet. These operations included the Miller, Smith and
Pope farms, on the west side of the river, and the Rice, Nicklin,
Martin and Harmon, on the east side, all second-sand territory. North
of the Cochran and the Hoover work was pushed actively. George H.
Bissell and Vance Stewart bored twelve or fifteen medium wells on
the Stewart farm of two-hundred acres, which the Cameron
Petroleum Company purchased in 1865 and Joseph Dale operated for
some years. It lies below the lower bridge, opposite the Bleakley
tract, from which a light production is still derived. Above the Stewart
are the Fuller and the Chambers farms, the latter extending to the
Allegheny-Valley depot. Scores of eager operators thronged the
streets of Franklin and drilled along the Allegheny. Joseph Powley
and Charles Cowgill entered the lists in the Cranberry district. Henry
M. Wilson and George Piagett veered into the township and sank a
bevy of dry-holes to vary the monotony. That was a horse on Wilson,
but he got ahead of the game by a deal that won him the nicest
territory on Horse Creek. Stirling Bonsall and Colonel Lewis—they’re
dead now—were in the thickest of the fray, with Captain Goddard,
Philip Montgomery, Boyd, Roberts, Foster, Brown, Murphy and many

more whom old-timers remember pleasantly. Thomas King, whole-
souled, genial “Tom”—no squarer man e’er owned a well or handled
oil-certificates—and Captain Griffith were “a good pair to draw to.”
King has “crossed over,” as have most of the kindred spirits that
dispelled the gloom in the sixties.
Colonel W. T. Pelton, nephew of Samuel J. Tilden, participated in
the scenes of that exciting period. He lived at Franklin and drilled
wells on French Creek. He was a royal entertainer, shrewd in
business, finely educated and polished in manner and address. He
and his wife—a lovely and accomplished woman—were fond of
society and gained hosts of friends. They boarded at the United-
States Hotel, where Mrs. Pelton died suddenly. This affliction led
Colonel Pelton to sell his oil-properties and abandon the oil-regions.
Returning to New York, when next he came into view as the active
agent of his uncle in the secret negotiations that grew out of the
election of 1876, it was with a national fame. His death in 1880
closed a busy, promising career.
In the spring of 1864 a young man, black-haired, dark-eyed, an
Apollo in form and strikingly handsome, arrived at Franklin and
engaged rooms at Mrs. Webber’s, on Buffalo street. The stranger
had money, wore good clothes and presented a letter of introduction
to Joseph H. Simonds, dealer in real-estate, oil-wells and leases. He
looked around a few days and concluded to invest in sixty acres of
the Fuller farm, Cranberry township, fronting on the Allegheny river.
The block was sliced off the north end of the farm, a short distance
below the upper bridge and the Valley station. Mr. Simonds
consented to be a partner in the transaction. The transfer was
effected, the deed recorded and a well started. It was situated on
the hill, had twenty feet of second-sand and pumped twenty barrels
a day. The owner drilled two others on the bluff, the three yielding
twenty barrels for months. The ranks of the oil-producers had
received an addition in the person of—John Wilkes Booth.
The firm prospered, each of the members speculating and trading
individually. M. J. Colman, a capital fellow, was interested with one
or both in various deals. Men generally liked Booth and women
admired him immensely. His lustrous orbs, “twin-windows of the

soul,” could look so sad and pensive as to awaken the tenderest pity,
or fascinate like “the glittering eye” of the Ancient Mariner or the
gaze of the basilisk. “Trilby” had not come to light, or he might have
enacted the hypnotic role of Svengali. His moods were variable and
uncertain. At times he seemed morose and petulant, tired of
everybody and “unsocial as a clam.” Again he would court society,
attend parties, dance, recite and be “the life of the company.” He
belonged to a select circle that exchanged visits with a coterie of
young folks in Oil City. A Confederate sympathizer and an enemy of
the government, his closest intimates were staunch Republicans and
loyal citizens. William J. Wallis, the veteran actor who died in
December of 1895, in a Philadelphia theater slapped him on the
mouth for calling President Lincoln a foul name. Booth’s acting, while
inferior to his brother Edwin’s, evinced much dramatic power. He
controlled his voice admirably, his movements were graceful and he
spoke distinctly, as Franklinites whom he sometimes favored with a
reading can testify.
JOSEPH H. SIMONDS.

J. WILKES BOOTH.

MOSES J. COLMAN.
One morning in April, 1865, he left Franklin, telling Mr. Simonds he
was going east for a few days. He carried a satchel, which indicated
that he did not expect his stay to be prolonged indefinitely. His
wardrobe, books and papers remained in his room. Nothing was
heard of him until the crime of the century stilled all hearts and the

wires flashed the horrible news of Abraham Lincoln’s assassination.
The excitement in Franklin, the murderer’s latest home, was intense.
Crowds gathered to learn the dread particulars and discuss Booth’s
conduct and utterances. Not a word or act previous to his departure
pointed to deliberate preparation for the frightful deed that plunged
the nation in grief. That he contemplated it before leaving Franklin
the weight of evidence tended to disprove. He made no attempt to
sell any of his property, to convert his lands and wells into cash, to
settle his partnership accounts or to pack his effects. He had money
in the bank, wells bringing a good income and important business
pending. All these things went to show that, if not a sudden impulse,
the killing of Lincoln was prompted by some occurrence in
Washington that fired the passionate nature John Wilkes Booth
inherited from his father. The world is familiar with the closing
chapters of the dark tragedy—the assassin’s flight, the pursuit into
Virginia, the burning barn, Sergeant Corbett’s fatal bullet, the
pathetic death-scene on the Garrett porch and the last message, just
as the dawn was breaking on the glassy eyes that opened feebly for
a moment: “Tell my mother I died for my country. I did what I
thought was best.”
The wells and the land on the river were held by Booth’s heirs until
1869, when the tract changed hands. The farm is producing no oil
and the Simonds-Booth wells have disappeared. Had he not intended
to return to Franklin, Booth would certainly have disposed of these
interests and given the proceeds to his mother. “Joe” Simonds
removed to Bradford to keep books for Whitney & Wheeler, bankers
and oil-operators, and died there years ago. He was an expert
accountant, quick, accurate and neat in his work and most fastidious
in his attire. A blot on his paper, a figure not exactly formed, a line
one hair-breath crooked, a spot on his linen or a speck of dust on his
coat was simply intolerable. He was correct in language and
deportment and honorable in his dealings. Colman continued his oil-
operations, in company with W. R. Crawford, a real-estate agency,
until the eighties. He married Miss Ella Hull, the finest vocalist
Franklin ever boasted, daughter of Captain S. A. Hull, and removed

to Boston. For years paralytic trouble has confined him to his home.
He is “one of nature’s nobleman.”
“French Kate,” the woman who aided Ben Hogan at Pithole and
followed him to Babylon and Parker, was a Confederate spy and
supposed to be very friendly with J. Wilkes Booth. Besides his oil-
interests at Franklin, the slayer of Abraham Lincoln owned a share in
the Homestead well at Pithole. A favorite legend tells how, by a
singular coincidence, which produced a sensation, the well was
burned on the evening of the President’s assassination. It caught fire
about the same instant the fatal bullet was fired in Ford’s Theater
and tanks of burning oil enveloped Pithole in a dense smoke when
the news of the tragedy flashed over the trembling wires. The
Homestead well was not down until Lincoln had been dead seven
weeks, Pithole had no existence and there were no blazing tanks;
otherwise the legend is correct. Two weeks before his appalling
crime Booth was one of a number of passengers on the scow doing
duty as a ferry-boat across the Allegheny, after the Franklin bridge
had burned. The day was damp and the water very cold. Some
inhuman whelp threw a fine setter into the river. The poor beast
swam to the rear of the scow and Booth pulled him on board. He
caressed the dog and bitterly denounced the fellow who could treat
a dumb animal so cruelly. At another time he knocked down a
cowardly ruffian for beating a horse that was unable to pull a heavy
load out of a mud-hole. He has been known to shelter stray kittens,
to buy them milk and induce his landlady to care for them until they
could be provided with a home. Truly his was a contradictory nature.
He sympathized with horses, dogs and cats, yet robbed the nation of
its illustrious chief and plunged mankind into mourning. To newsboys
Booth was always liberal, not infrequently handing a dollar for a
paper and saying: “No change; buy something useful with the
money.” The first time he went to the Methodist Sunday-school, with
“Joe” Simonds, he asked and answered questions and put a ten-
dollar bill in the collection-box.
Over the hills to the interior of the townships developments
spread. Bredinsburg, Milton and Tarkiln loomed up in Cranberry,
where Taylor & Torrey, S. P. McCalmont, Jacob Sheasley, B. W. Bredin

and E. W. Echols have sugar-plums. In Sandycreek, between Franklin
and Foster, Angell & Prentice brought Bully Hill and Mount Hope to
the front. The biggest well in the package was a two-hundred
barreler on Mount Hope, which created a mount of hopes that were
not fully realized. George V. Forman counted out one-hundred-and-
fifty-thousand dollars for the Mount Hope corner. The territory lasted
well and averaged fairly. Bully Hill merited its somewhat slangy title.
Dr. C. D. Galbraith, George R. Sheasley and Mattern & Son are
among its present operators. Angell and Prentice parted company,
each to engage in opening up the Butler region. Prentice, Crawford,
Barbour & Co. did not let the grass grow under their feet. They
“knew a good thing at sight” and pumped tens-of-thousands of
barrels of oil from the country south of Franklin. The firm was
notable in the seventies. Considerable drilling was done at Polk,
where the state is providing a half-million-dollar Home for Feeble-
Minded Children, and in the latitude of Utica, with about enough oil
to be an aggravation. The Shippen wells, a mile north of the county
poor-house, have produced for thirty years. West of them, on the
Russell farm, the Twin wells, joined as tightly as the derricks could
be placed, pumped for years. This was the verge of productive
territory, test wells on the lands of William Sanders, William Bean, A.
Reynolds, John McKenzie, Alexander Frazier and W. Booth, clear to
Cooperstown, finding a trifle of sand and scarcely a vestige of oil.
The Raymonds, S. Ramage, John J. Doyle and Daniel Grimm had a
very tidy offshoot at Raymilton. On this wise lubricating and second-
sand oils were revealed for the benefit of mankind generally. The fly
in the ointment was the clerical crank who wrote to President Lincoln
to demand that the producing of heavy-oil be stopped peremptorily,
as it had been stored in the ground to grease the axletree of the
earth in its diurnal revolution! This communication reminded Lincoln
of a “little story,” which he fired at the fellow with such effect that
the candidate for a strait-jacket was perpetually squelched.

JOHN P. CRAWFORD.
ANGELL & PRENTICE’S WELLS BELOW FRANKLIN IN 1873.
Hon. William Reid Crawford, a member
of the firm of Prentice, Crawford, Barbour
& Co., lives in Franklin. His parents were
early settlers in north-western
Pennsylvania. Alexander Grant, his
maternal grandfather, built the first stone-
house in Lancaster county, removed to
Butler county and located finally in
Armstrong county, where he died sixty-five
years ago. In 1854 William R. and four of
his brothers went to California and spent
some time mining gold. Upon his return
he settled on a farm in Scrubgrass
township, Venango county, of which
section the Crawfords had been prominent
citizens from the beginning of its history.
Removing to Franklin in 1865, Mr. Crawford engaged actively in the
production of petroleum, operating extensively in various portions of

the oil-regions for twenty years. He acquired a high reputation for
enterprise and integrity, was twice a city-councillor, served three
terms as mayor, was long president of the school-board, was elected
sheriff in 1887 and State-Senator in 1890. Untiring fidelity to the
interests of the people and uncompromising hostility to whatever he
believed detrimental to the general welfare distinguished his public
career. Genial and kindly to all, the friend of humanity and
benefactor of the poor, no man stands better in popular estimation
or is more deserving of confidence and respect. His friends could not
be crowded into the Coliseum without bulging out the walls.
Ebenezer Crawford, brother of William R., died at Emlenton in August
of 1897, on his seventy-sixth birthday. John P. Crawford, another
brother, who made the California trip in 1849, still resides in the
southern end of the county and is engaged in oil-operations. E. G.
Crawford, a nephew, twice prothonotary of Venango and universally
liked, passed away last June. His cousin, C. J. Crawford, a first-class
man anywhere and everywhere, served as register and recorder with
credit and ability. The Crawfords “are all right.”
For money may come and money may go,
But a good name stays to the end of the show.
Captain John K. Barbour, a man of imposing presence and
admirable qualities, removed to Philadelphia after the dissolution of
the firm. The Standard Oil-Company gave him charge of the right-of-
way department of its pipe-line service and he returned to Franklin.
Two years ago, during a business visit to Ohio, he died unexpectedly,
to the deep regret of the entire community. S. A. Wheeler operated
largely in the Bradford field and organized the Tuna-Valley Bank of
Whitney & Wheeler. For a dozen years he has resided at Toledo, his
early home. Like Captain Barbour, “Fred,” as he was commonly
called, had an exhaustless mine of bright stories and a liberal share
of the elements of popularity. One afternoon in 1875, three days
before the fire that wiped out the town, a party of us chanced to
meet at St. Joe, Butler county, then the centre of oil-developments.
An itinerating artist had his car moored opposite the drug-store.
Somebody proposed to have a group-picture. The motion carried

unanimously and a toss-up decided that L. H. Smith was to foot the
bill. The photographer brought out his camera, positions were taken
on the store-platform and the pictures were mailed an hour ahead of
the blaze that destroyed most of the buildings and compelled the
artist to hustle off his car on the double-quick. Samuel R. Reed, at
the extreme right, operated in the Clarion field. He had a hardware-
store in company with the late Dr. Durrant and his home is in
Franklin. James Orr, between whom and Reed a telegraph-pole is
seen, was connected with the Central Hotel at Petrolia and later was
a broker in the Producers’ Exchange at Bradford. On the step is
Thomas McLaughlin, now oil-buyer at Lima, once captain of a
talented base-ball club at Oil City and an active oil-broker. Back of
him is “Fred” Wheeler, with Captain Barbour on his right and L. H.
Smith sitting comfortably in front. Mr. Smith figured largely at
Pithole, operated satisfactorily around Petrolia and removed years
ago to New York. Cast in a giant mould, he weighs three-hundred
pounds and does credit to the illustrious legions of Smiths. He is a
millionaire and has an office over the Seaboard Bank, at the lower
end of Broadway. Joseph Seep, the king-bee of good fellows, sits
besides Smith. Pratt S. Crosby, formerly a jolly broker at Parker and
Oil City, stands behind Seep. Next him is “Tom” King, who has “gone
to the land of the leal,” J. J. McLaurin ending the row. James Amm,
who went from an Oil-city clerkship to coin a fortune at Bradford—a
street bears his name—sits on the platform. Every man, woman,
child and baby near Oil City knew and admired “Jamie” Amm, who is
now enjoying his wealth in Buffalo. Two out of the eleven in the
group have “passed beyond the last scene” and the other nine are
scattered widely.
“Friend after friend departs,
Who hath not lost a friend?”
Frederic Prentice, one of the pluckiest operators ever known in
petroleum-annals, was the first white child born on the site of
Toledo, when Indians were the neighbors of the pioneers of Northern
Ohio. His father left a fine estate, which the son increased greatly by
extensive lumbering, in which he employed three-thousand men.

GROUP AT ST. JOE, BUTLER COUNTY, IN 1874.
Losses in the panic of
1857 retired him from
the business. He
retrieved his fortune and
paid his creditors their
claims in full, with ten
per cent. interest, an act
indicative of his sterling
character. Reading in a
newspaper about the
Drake well, he decided to
see for himself whether
the story was fast colors.
Journeying to Venango
county by way of
Pittsburg, he met and engaged William Reed to accompany him.
Reed had worked at the Tarentum salt-wells and knew a thing or two
about artesian-boring. The two arrived at Franklin on the afternoon
of the day Evans’s well turned the settlement topsy-turvy. Next
morning Prentice offered Evans forty-thousand dollars for a
controlling interest in the well, one-fourth down and the balance in
thirty, sixty and ninety days. Evans declining to sell, the Toledo visitor
bought from Martin & Epley an acre of ground on the north bank of
French Creek, at the base of the hill, and contracted with Reed to
“kick down” a well, the third in the district. Prentice and Reed
tramped over the country for days, locating oil-deposits by means of
the witch-hazel, which the Tarentumite handled skillfully. This was a
forked stick, which it was claimed turned in the hands of the holder
at spots where oil existed. Various causes delayed the completion of
the well, which at last proved disappointingly small. Meanwhile Mr.
Prentice leased the Neeley farm, two miles up the Allegheny, in
Cranberry township, and bored several paying wells. A railroad
station on the tract is named after him and R. G. Lamberton has
converted the property into a first-class stock-farm. Favorable reports
from Little Kanawha River took him to West Virginia, where he leased
and purchased immense blocks of land. Among them was the Oil-

FREDERIC PRENTICE.
Springs tract, on the Hughes River, from which oil had been skimmed
for generations. Two of his wells on the Kanawha yielded six-hundred
barrels a day, which had to be stored in ponds or lakes for want of
tankage. Confederate raiders burned the wells, oil and machinery
and drove off the workmen, putting an extinguisher on operations
until the Grant-Lee episode beneath the apple-tree at Appomattox.
Assuming that the general direction of profitable developments
would be north-east and south-west, Mr. Prentice surveyed a line
from Venango county through West Virginia, Kentucky and
Tennessee. This idea, really the foundation of “the belt theory,” he
spent thousands of dollars to establish. Personal investigation and
careful surveys confirmed his opinion, which was based upon
observations in the Pennsylvania fields. The line run thirty years ago
touched numerous “springs” and “surface shows” and recent tests
prove its remarkable accuracy. On this theory he drilled at Mount
Hope and Foster, opening a section that has produced several-million
barrels of oil. C. D. Angell applied the principle in Clarion and Butler
counties, mapping out the probable course of the “belt” and leasing
much prolific territory. His success led others to adopt the same plan,
developing a number of pools in four states, although nature’s lines
are seldom straight and the oil-bearing strata are deposited in curves
and beds at irregular intervals.
In company with W. W. Clark of New
York, to whom he had traded a portion of
his West-Virginia lands, Mr. Prentice
secured a quarter-interest in the Tarr
farm, on Oil Creek, shortly before the
sinking of the Phillips well, and began
shipping oil to New York. They paid three
dollars apiece for barrels, four dollars a
barrel for hauling to the railroad and
enormous freights to the east. The price
dropping below the cost of freights and
barrels, the firm dug acres of pits to put
tanks under ground, covering them with
planks and earth to prevent evaporation.

Traces of these storage-vats remain on the east bank of Oil Creek.
Crude fell to twenty-five cents a barrel at the wells and the outlook
was discouraging. Clark & Prentice stopped drilling and turned their
attention to finding a market. They constructed neat wooden
packages that would hold two cans of refined-oil, two oil-lamps and
a dozen chimneys and sent one to each United-States Consul in
Europe. Orders soon rushed in from foreign countries, especially
Germany, France and England, stimulating the erection of refineries
and creating a large export-trade. Clark & Summer, who also owned
an interest in the Tarr farm, built the Standard Refinery at Pittsburg
and agreed to take from Clark & Prentice one-hundred-thousand
barrels of crude at a dollar a barrel, to be delivered as required
during the year. Before the delivery of the first twenty-five-thousand
barrels the price climbed to one-fifty and to six dollars before the
completion of the contract, which was carried out to the letter. The
advance continued to fourteen dollars a barrel, lasting only one day
at this figure. These were vivifying days in oleaginous circles, never
to be repeated while Chronos wields his trusty blade.
When crude reached two dollars Mr. Prentice bought the
Washington-McClintock farm, on which Petroleum Centre was
afterwards located, for three-hundred-thousand dollars. Five New-
Yorkers, one of them the president of the Shoe and Leather Bank
and another the proprietor of the Brevoort House, advanced fifty-
thousand dollars for the first payment. Within sixty days Prentice
sold three-quarters of his interest for nine-hundred-thousand dollars
and organized the Central Petroleum Oil-Company, with a capital of
five-millions! Wishing to repay the New-York loan, the Brevoort
landlord desired him to retain his share of the money and invest it as
he pleased. For his ten-thousand dollars mine host received eighty-
thousand in six months, a return that leaves government-bond
syndicates and Cripple-Creek speculations out in the latitude of
Nansen’s north-pole. The company netted fifty-thousand dollars a
month in dividends for years and lessees cleared three or four
millions from their operations on the farm. Greenbacks circulated like
waste-paper, Jules Verne’s fancies were surpassed constantly by
actual occurrences and everybody had money to burn.

Prentice and his associates purchased many tracts along Oil Creek,
including the lands where Oil City stands and the Blood farm of five-
hundred acres. In the Butler district he drilled hundreds of wells and
built the Relief Pipe-Line. Organizing The Producers’ Consolidated
Land-and-Petroleum-Company, with a capital of two-and-a-half
millions, he managed it efficiently and had a prominent part in the
Bradford development. Boston capitalists paid in twelve-hundred-
thousand dollars, Prentice keeping a share in his oil-properties
representing thirteen-hundred-thousand more. The company is now
controlled by the Standard, with L. B. Lockhart as superintendent. Its
indefatigable founder also organized the Boston Oil Company to
operate in Kentucky and Tennessee, put down oil-wells in Peru and
gas-wells in West Virginia, produced and piped thousands of barrels
of crude daily and was a vital force in petroleum-affairs for eighteen
years. The confidence and esteem of his compatriots were attested
by his unanimous election to the presidency of the Oilmen’s League,
a secret-society formed to resist the proposed encroachments of the
South-Improvement Company. The League accomplished its mission
and then quietly melted out of existence.
Since 1877 Mr. Prentice has devoted his attention chiefly to
lumbering in West Virginia and to his brown-stone quarries at
Ashland, Wisconsin. The death of his son, Frederick A., by accidental
shooting, was a sad bereavement to the aged father. His suits to get
possession of the site of Duluth, the city of Proctor Knott’s
impassioned eulogy, included in a huge grant of land deeded to him
by the Indians, were scarcely less famous than Mrs. Gaines’s
protracted litigation to recover a slice of New Orleans. The claim
involved the title to property valued at twelve-millions of dollars.
From his Ashland quarries the owner took out a monolith, designed
for the Columbian Exposition in 1893, forty yards long and ten feet
square at the base. Beside this monster stone Cleopatra’s Needle,
disintegrating in Central Park, Pompey’s Pillar and the biggest blocks
in the pyramids are Tom-Thumb pigmies. At seventy-four Mr.
Prentice, foremost in energy and enterprise, retains much of his
youthful vigor. Earnest and sincere, a master of business, his word as

CYRUS D. ANGELL.
good as gold, Frederic Prentice holds an honored place in the ranks
of representative oil-producers, “nobles of nature’s own creating.”
A native of Chautauqua county, N. Y.,
where he was born in 1826, Cyrus D.
Angell received a liberal education, served
as School-Commissioner and engaged in
mercantile pursuits at Forestville. Forced
through treachery and the monetary
stringency of the times to compromise
with his creditors, he recovered his
financial standing and paid every cent of
his indebtedness, principal and interest. In
1867 he came to the oil-regions with a
loan of one-thousand dollars and
purchased an interest in property at
Petroleum Centre that paid handsomely.
Prior to this, in connection with Buffalo
capitalists, he had bought Belle Island, in the Allegheny River at
Scrubgrass, upon which soon after his arrival he drilled three wells
that averaged one-hundred barrels each for two years, netting the
owners over two-hundred-thousand dollars. Operations below
Franklin, in company with Frederic Prentice, also proved highly
profitable. His observations of the course of developments along Oil
Creek and the Allegheny led Mr. Angell to the conclusion that
petroleum would be found in “belts” or regular lines. He adopted the
theory that two “belts” existed, one running from Petroleum Centre
to Scrubgrass and the other from St. Petersburg through Butler
county. Satisfied of the correctness of this view, he leased or
purchased all the lands within the probable boundaries of the “belt”
from Foster to Belle Island, a distance of six miles. The result
justified his expectations, ninety per cent. of the wells yielding
abundantly. With “the belt theory,” which he followed up with equal
success farther south, Mr. Angell’s name is linked indissolubly. His
researches enriched him and were of vast benefit to the producers
generally. He did much to extend the Butler region, drilling far ahead
of tested territory. The town of Angelica owed its creation to his

fortunate operations in the neighborhood, conducted on a
comprehensive scale. Reverses could not crush his manly spirit. He
did a large real-estate business at Bradford for some years, opening
an office at Pittsburg when the Washington field began to loom up.
Failing health compelling him to seek relief in foreign travel, last year
he went to Mexico and Europe to recuperate. Mr. Angell is endowed
with boundless energy, fine intellectual powers and rare social
acquirements. During his career in Oildom he was an excellent
sample of the courageous, unconquerable men who have made
petroleum the commercial wonder of the world.
An old couple in Cranberry township, who eked out a scanty living
on a rocky farm near the river, sold their land for sixty-thousand
dollars at the highest pitch of the oil-excitement around Foster. This
was more money than the pair had ever before seen, much less
expected to handle and own. It was paid in bank-notes at noon and
the log-house was to be vacated next day. Towards evening the poor
old woman burst into tears and insisted that her husband should
give back the money to the man that “wanted to rob them of their
home.” She was inconsolable, declaring they would be “turned out to
starve, without a roof to cover them.” The idea that sixty-thousand
dollars would buy an ideal home brought no comfort to the simple-
minded creature, whose hopes and ambitions were confined to the
lowly abode that had sheltered her for a half-century. A promise to
settle near her brother in Ohio reconciled her somewhat, but it
almost broke her faithful heart to leave a spot endeared by many
tender associations. John Howard Payne, himself a homeless
wanderer, whose song has been sung in every tongue and echoed in
every soul, jingled by innumerable hand-organs and played by the
masters of music, was right:
“Be it ever so humble, there’s no place like home.”
The refusal of his wife to sign the deed conveying the property
enabled a wealthy Franklinite to gather a heap of money. The tract
was rough and unproductive and the owner proposed to accept for it
the small sum offered by a neighboring farmer, who wanted more

pasture for his cattle. For the first time in her life the wife declined to
sign a paper at her husband’s request, saying she had a notion the
farm would be valuable some day. The purchaser refused to take it
subject to a dower and the land lay idle. At length oil-developments
indicated that the “belt” ran through the farm. Scores of wells
yielded freely, netting the land-owner a fortune and convincing him
that womanly intuition is a sure winner.
A citizen of Franklin, noted for his conscientiousness and liberality,
was interested in a test-well at the beginning of the Scrubgrass
development. He vowed to set aside one-fourth of his portion of the
output of the well “for the Lord,” as he expressed it. To the delight of
the owners, who thought the venture hazardous, the well showed for
a hundred barrels when the tubing was put in. On his way back from
the scene the Franklin gentleman did a little figuring, which proved
that the Lord’s percentage of the oil might foot up fifty dollars a day.
This was a good deal of money for religious purposes. The maker of
the vow reflected that the Lord could get along without so much
cash and he decided to clip the one-fourth down to one-tenth,
arguing that the latter was the scripture limit. Talking it over with his
wife, she advised him to stick to his original determination and not
trifle with the Lord. The husband took his own way, as husbands are
prone to do, and revisited the well next day. Something had gone
wrong with the working-valve, the tubing had to be drawn out and
the well never pumped a barrel of oil! The disappointed operator
concluded, as he charged two thousand dollars to his profit-and-loss
account, that it was not the Lord who came out at the small end of
the horn in the transaction.
Rev. Clarence A. Adams, the eloquent ex-pastor of the First Baptist
Church at Franklin, is the lucky owner of a patch of paying territory
at Raymilton. Recently he finished a well which pumped considerable
salt-water with the oil. Contrary to Cavendish and the ordinary
custom, another operator drilled very close to the boundary of the
Adams lease and torpedoed the well heavily. Instead of sucking the
oil from the preacher’s nice pumper, the new well took away most of
the salt-water and doubled the production of petroleum! Commonly
it would seem rather mean to rob a Baptist minister of water, but in

REV. C. A. ADAMS, D.D. REV. EZRA F. CRANE, D.D.
this case Dr.
Adams is
perfectly
resigned to the
loss of aqueous
fluid and gain of
dollar-fifty
crude. A
profound
student of
Shakespeare,
Browning and
the Bible, a
brilliant lecturer
and master of
pulpit-oratory, may he also stand on a lofty rung of the greasian
ladder and attain the goodly age of Franklin’s “grand old man,” Rev.
Dr. Crane. This “father in Israel,” whose death in February of 1896
the whole community mourned, left a record of devoted service as a
physician and clergyman for over sixty years that has seldom been
equaled. He healed the sick, smoothed the pillow of the dying,
relieved the distressed, reclaimed the erring, comforted the
bereaved, turned the faces of the straying Zionward and found the
passage to the tomb “a gentle wafting to immortal life.” Let his
memory be kept green.
“Though old, he still retained
His manly sense and energy of mind.
Virtuous and wise he was, but not severe,
For he remembered that he once was young;
His kindly presence checked no decent joy.
Him e’en the dissolute admired. Can he be dead
Whose spiritual influence is upon his kind?”
The late Thomas McDonough, a loyal-hearted son of the Emerald
Isle, was also an energetic operator in the lubricating region. He had
an abundance of rollicking wit, “the pupil of the soul’s clear eye,” and

an unfailing supply of the drollest stories. Desiring to lease a farm in
Sandy-Creek township, supposed to be squarely “on the belt,” he
started at daybreak to interview the owner, feeling sure his mission
would succeed. An unexpected sight presented itself through the
open door, as the visitor stepped upon the porch of the dwelling. The
farmer’s wife was setting the table for breakfast and Frederic
Prentice was folding a paper carefully. McDonough realized in a
twinkling that Prentice had secured the lease and his trip was
fruitless. “I am looking for John Smith” he stammered, as the farmer
invited him to enter, and beat a hasty retreat. For years his friends
rallied the Colonel on his search and would ask with becoming
solemnity whether he had discovered John Smith. The last time we
met in Philadelphia this incident was revived and the query repeated
jocularly. The jovial McDonough died in 1894. It is safe to assume
that he will easily find numerous John Smiths in the land of
perpetual reunion. One day he told a story in an office on Thirteenth
street, Franklin, which tickled the hearers immensely. A full-fledged
African, who had been sweeping the back-room, broke into a
tumultuous laugh. At that moment a small boy was riding a donkey
directly in front of the premises. The jackass heard the peculiar
laugh and elevated his capacious ears more fully to take in the
complete volume of sound. He must have thought the melody
familiar and believed he had stumbled upon a relative. Despite the
frantic exertions of the boy, the donkey rushed towards the building
whence the boisterous guffaw proceeded, shoved his head inside the
door and launched a terrific bray. The bystanders were convulsed at
this evidence of mistaken identity, which the jolly story-teller
frequently rehearsed for the delectation of his hosts of friends.
Looking over the Milton diggings one July day, Col. McDonough
met an amateur-operator who was superintending the removal of a
wooden-tank from a position beside his first and only well. A
discussion started regarding the combustibility of the thick sediment
collected on the bottom of the tank. The amateur maintained the
stuff would not burn and McDonough laughingly replied, “Well, just
try it and see!” The fellow lighted a match and applied it to the viscid
mass before McDonough could interfere, saying with a grin that he

THOMAS M’DONOUGH.
proposed to wait patiently for the result.
He didn’t have to wait “until Orcus would
freeze over and the boys play shinny on
the ice.” In the ninetieth fraction of a
second the deposit blazed with intense
enthusiasm, quickly enveloping the well-
rig and the surroundings in flames. Clouds
of smoke filled the air, suggesting fancies
of Pittsburg or Sheol. Charred fragments
of the derrick, engine-house and tank,
with an acre of blackened territory over
which the burning sediment had spread,
demonstrated that the amateur’s idea had
been decidedly at fault. The experiment convinced him as
searchingly as a Roentgen ray that McDonough had the right side of
the argument. “If the ‘b. s.’ had been as green as the blamed fool, it
wouldn’t have burned,” was the Colonel’s appropriate comment.
Miss Lizzie Raymond, daughter of the pioneer who founded
Raymilton and erected the first grist-mill at Utica, has long taught
the infant-class of the Presbyterian Sunday-school at Franklin. Once
the lesson was about the wise and the foolish virgins, the good
teacher explaining the subject in a style adapted to the juvenile
mind. A cute little tot, impressed by the sad plight of the virgins who
had no oil in their lamps, innocently inquired: “Miss ’Aymond, tan’t
oo tell ’em dirls to turn to our house an’ my papa ’ll div’ ’em oil f’um
his wells?” Heaven bless the children that come as sunbeams to
lighten our pathway, to teach us lessons of unselfishness and
prevent the rough world from turning our hearts as hard as the mill-
stone.
Judge Trunkey, who presided over the Venango court a dozen
years and was then elected to the Supreme Bench, was hearing a
case of desertion. An Oil-City lawyer, proud of his glossy black beard,
represented the forsaken wife, a comely young woman from
Petroleum Centre, who dandled a bright baby of twenty months on
her knee. Mother and baby formed a pretty picture and the lawyer
took full advantage of it in his closing appeal to the jury. At a brilliant

climax he turned to his client and said: “Let me have the child!” He
was raising it to his arms, to hold before the men in the box and
describe the heinous meanness of the wretch who could leave such
beauty and innocence to starve. The baby spoiled the fun by
springing up, clutching the attorney’s beard and screaming: “Oh,
papa!” The audience fairly shrieked. Judge Trunkey laughed until the
tears flowed and it was five minutes before order could be restored.
That ended the oratory and the jury salted the defendant
handsomely. Hon. James S. Connelly, an Associate Judge, who now
resides in Philadelphia and enjoys his well-earned fortune, was also
on the bench at the moment. Judge Trunkey, one of the purest,
noblest men and greatest jurists that ever shed lustre upon
Pennsylvania, passed to his reward six years ago.
In your wide peregrinations from the poles to the equator,
Should you hear some ignoramus—let out of his incubator—
Say the heavy-oil of Franklin is not earth’s best lubricator,
Do as did renown’d Tom Corwin, the great Buckeye legislator,
When a jabberwock in Congress sought to brand him as a traitor,
Just “deny the allegation and defy the allegator!”
MILLER & SIBLEY’S PROSPECT-HILL STOCK FARM FRANKLIN, PA.

KEEPING STEP.
The Shasta was Karns City’s first well.
Missouri has two wells producing oil.
North Dakota has traces of natural-gas.
Ninety wells in Japan pump four-hundred barrels.
Elk City, in the Clarion field, once had two-thousand population.
The Rob Roy well, at Karns City, has produced a quarter-million
barrels of oil.
Alaska-oil is cousin of asphalt-pitch, very heavy, and thick as New-
Orleans molasses in midwinter.
Wade Hampton, postmaster of Pittsburg, and cousin of Governor
Wade Hampton, organized one of the first petroleum companies in
the United States.
General Herman Haupt, of Philadelphia, now eighty-one years old,
surveyed the route and constructed the first pipe-line across
Pennsylvania.
Robert Nevin, founder of the Pittsburg Times, drilled a dry-hole
four-hundred feet, ten miles west of Greensburg, in 1858, a year
before Drake’s successful experiment in Oil Creek.
The Powell Oil-Company, superintended by Col. A. C. Ferris, still a
resident of New York, paid fifty-thousand dollars in cash for the Shirk
farm, half way between Franklin and Oil City, drilled a dry-hole and
abandoned the property.
The gentle wife who seeks your faults to cover
You don’t deserve; prize naught on earth above her;
Keep step and be through life her faithful lover.
The new town of Guffey, the liveliest in Colorado, thirty miles from
Cripple Creek, is fitly named in honor of James M. Guffey, the
successful Pennsylvania oil-producer and political leader, who has big
mining interests in that section.

The Fonner pool, Greene county, was the oil-sensation of 1897 in
Pennsylvania. The Fonner well, struck in March, and territory around
it sold for two-hundred-thousand dollars. Elk Fork wore the West-
Virginia belt, Peru took the Hoosier biscuit and Lucas county the
Buckeye premium.
Say, boys, seein’ how fast th’ ranks iz thinnin’—
Th’ way thar droppin’ out sets my head spinnin’—
An’ knownin’ ez how death may take an innin’
An’ clean knock out our underpinnin’,
I kalkilate we oughter swar off sinnin’,
Jes’ quit fer keeps our dog-gon’ chinnin’,
Start in th’ narrer road fer a beginning’,
An’ so strike oil in Heav’n fer a sure winnin’
When up the golden-stairs we goes a-shinnin’.
When the biggest well in Indiana flowed oil fifty feet above the
derrick, at Van Buren, a local paper noted the effect thus: “The strike
has given the town a tremendous boom. Several real-estate offices
have opened and the town-council has raised the license for faro-
banks from five dollars a year to twelve dollars.” At this rate Van
Buren ought soon to be in the van.

JOHN VANAUSDALL. WM. PHILLIPS.
GEO. K. ANDERSON.
F. S. TARBELL. F. W. ANDREWS.
ORIGINAL D. W. KENNEY’S ALLEMAGOOZELUM-CITY
WELL N
o
2.
CAPT. WM. HASSON. JOHN P. ZANE.
HENRY R. ROUSE.

VII.
THE VALLEY OF PETROLEUM.
Wonderful Scenes on Oil Creek—Mud and Grease Galore—Rise and Fall of Phenomenal
Towns—Shaffer , Pioneer and Petroleum Centre—Fortune’s Queer Vagaries—Wells
Flowing Thousands of Barrels —Sherman , Delamater and “Coal-Oil Johnnie ”—From
Penury to Riches and Back—Recitals That Discount Fairy-Tales.
“I pity the man who can travel from Dan to Beersheba and cry, ‘’Tis all barren.’”—
Sterne.
“This beginning part is not made out of anybody’s head; it’s real.”—Dickens.
“Some ships come into port that are not steered.”—Seneca.
“God has placed in his great bank—mother earth—untold wealth and many a poor
man’s check has been honored here for large amounts of oil.”—T. S. Scoville,
A. D. 1861.
“Ain’t that well spittin’ oil?”—Small Boy, A. D. 1863.
“Wonderful, most wonderful, marvelous, most marvelous, are the stories told of
the oil-region. It is another California.”—John W. Forney, A. D. 1863.
“Derricks peered up behind the houses of Oil City, like dismounted steeples, and
oil was pumping in the back-yards.”—London Post, A. D. 1865.
“From this place and from this day henceforth commences a new era.”—Goethe.
“The chandelier drives off with its splendor the darkness of night.”—Henry
Stanton.
“The onlookers were struck dumb with astonishment.”—Charles Kingsley.
“Either I will find a way or make one.”—Norman Proverb.
“I bid you look into the past as if it were a mirror.”—Terence.

orty-three farms of manifold shapes and
sizes lay along the stream from the
Drake well to the mouth of Oil Creek,
sixteen miles southward. For sixty
years the occupants of these tracts had
forced a bare subsistence from the
reluctant soil. “Content to live, to
propagate and die,” their requirements
and their resources were alike scanty.
They knew nothing of the artificial
necessities and extravagances of
fashionable life. To most of them the
great, busy, plodding world was a
sealed book, which they had neither the means nor the inclination to
unclasp. The world reciprocated by wagging in its customary groove,
blissfully unconscious of the scattered settlers on the banks of the
Allegheny’s tributary. A trip on a raft to Pittsburg, with the privilege
of walking back, was the limit of their journeyings from the hills and
rocks of Venango. Hunting, fishing and hauling saw-logs in winter
aided in replenishing the domestic larder. None imagined the
unproductive valley would become the cradle of an industry before
which cotton and coal and iron must “hide their diminished heads.”
No prophet had proclaimed that lands on Oil Creek would sell for
more than corner-lots in London or New York. Who could have
conceived that these bold cliffs and patches of clearing would enlist
ambitious mortals from every quarter of the globe in a mad race to
secure a foothold on the coveted acres? What seventh son of a
seventh son could foresee that a thousand dollars spent on the
Willard farm would yield innumerable millions? Who could predict
that a tiny stream of greenish fluid, pumped from a hole on an island
too insignificant to have a name, would swell into the vast ocean of
petroleum that is the miracle of the nineteenth century? Fortune has
played many pranks, but the queerest of them all were the vagaries
incidental to the petroleum-development on Oil Creek.
The Bissell, Griffin, Conley, two Stackpole, Pott, Shreve, two
Fleming, Henderson and Jones farms, comprising the four miles

between the Drake well and the Miller tract, were not especially
prolific. Traces of a hundred oil-pits, in some of which oak-trees had
grown to enormous size, are visible on the Bissell plot of eighty
acres. A large dam, used for pond-freshets, was located on Oliver
Stackpole’s farm. Two refineries of small capacity were built on the
Stackpole and Fletcher lands, where eighteen or twenty wells
produced moderately. The owner of a flowing well on the lower
Fleming farm, imitating the man who killed the goose that laid the
golden eggs, sought to increase its output by putting the tubing and
seed-bagging farther down. The well resented the interference,
refusing to yield another drop and pointing the obvious moral: “Let
well enough alone!” The Miller farm of four-hundred acres, on both
sides of the creek, was purchased in 1863 from Robert Miller by the
Indian Rock-Oil Company of New York. Now a railroad-station and
formerly the principal shipping-point for oil, refineries were started,
wells were drilled and the stirring town of Meredith blossomed for a
little space. The Lincoln well turned out sixty barrels a day, the
Boston fifty, the Bobtail forty, the Hemlock thirty and others from ten
to twenty-five, at an average depth of six-hundred feet. The
Barnsdall Oil-Company operated on the Miller and the Shreve farms,
drilling extensively on Hemlock Run, and George Bartlett ran the
Sunshine Oil-Works. The village, the refineries and the derricks have
disappeared as completely as Herculaneum or Sir John Franklin.
George Shaffer owned fifty acres below the Miller farm, divided by
Oil Creek into two blocks, one in Cherrytree township and the other
in Allegheny. Twenty-four wells, eight of them failures, were put
down on the flats and the abrupt hill bordering the eastern shore of
the stream. Samuel Downer’s Rangoon and three of Watson &
Brewer’s were the largest, ranking in the fifty-barrel list. In July of
1864 the Oil-Creek Railroad was finished to Shaffer farm, which
immediately became a station of great importance. From one house
and barn the place expanded in sixty days to a town of three-
thousand population. And such a town! Sixteen-hundred teams,
mainly employed to draw oil from the wells down the creek,
supported the stables, boarding-houses and hotels that sprang up in
a night. Every second door opened into a bar-room. The buildings

were “balloon frames,” constructed entirely of boards, erected in a
few hours and liable to collapse on the slightest pretext. Houses of
cards would be about as comfortable and substantial. Outdo
Hezekiah, by rolling back time’s dial thirty-one years, and in fancy
join the crowd headed for Shaffer six months after the advent of the
railway.
Start from Corry, “the city of stumps,” with the Downer refinery
and a jumble of houses thrown around the fields. Here the Atlantic &
Great-Western, the Philadelphia & Erie and the Oil-Creek Railroads
meet. The station will not shelter one-half the motley assemblage
bound for Oildom. “Mother Cary is plucking her geese” and snow-
flakes are dropping thickly. Speculators from the eastern cities,
westerners in quest of “a good thing,” men going to work at the
wells, capitalists and farmers, adventurers and drummers clamor for
tickets. It is the reverse of “an Adamless Eden,” for only three
women are to be seen. At last the train backs to the rickety depot
and a wild struggle commences. Scrambling for the elevated cars in
New York or Chicago is a feeble movement compared with this
frantic onslaught. Courtesy and chivalry are forgotten in the rush.
Men swarm upon the steps, clog the platforms, pack the baggage-
car, thrust the women aside, stick to the cowcatcher and clamber on
the roofs of the coaches. Over the roughest track on earth, which
winds and twists and skirts the creek most of the way, the train
rattles and jolts and pitches. The conductor’s job is no sinecure, as
he squeezes through the dense mass that leaves him without
sufficient elbow-room to “punch in the presence of the passenjare.”
Derricks—tall, gaunt skeletons, pickets of the advancing army—keep
solemn watch here and there, the number increasing as Titusville
comes in sight.
A hundred people get off and two-hundred manage somehow to
get on. Past the Drake well, past a forest of derricks, past steep cliffs
and tortuous ravines the engineer speeds the train. Did you ever
think what a weight of responsibility rests upon the brave fellow in
the locomotive-cab, whose clear eye looks straight along the track
and whose steady hand grasps the throttle? Should he relax his
vigilance or lose his nerve one moment, scores of lives might be the

fearful penalty. A short stop at Miller Farm, a whiff of refinery-smells
and in five minutes Shaffer is reached. The board-station is on the
right hand, landings on the left form a semi-circle hundreds of feet
in length, freight-cars jam the double track and warehouses dot the
bank. The flat-about thirty rods wide-contains the mushroom-town,
bristling with the undiluted essence of petroleum-activity. Three-
hundred teamsters are unloading barrels of oil from wagons dragged
by patient, abused horses and mules through miles of greasy, clayey
mud. Everything reeks with oil. It pervades the air, saturates clothes
and conversation, floats on the muddy scum and fills lungs and
nostrils with its peculiar odor. One cannot step a yard without sinking
knee-deep in deceptive mire that performs the office of a boot-jack if
given “a ghost of a show.” Christian’s Slough of Despond wasn’t a
circumstance to this adhesive paste, which engulfs unwary travelers
to their trouser-pockets and begets a dreadful craving for roads not
“Wholly unclassable,
Almost impassable,
Scarcely jackassable.”
The trip of thirty-five miles has shaken breakfast clear down to the
pilgrims’ boots. Out of the cars the hungry passengers tumble as
frantically as they had clambered in and break for the hotels and
restaurants. A dollar pays for a dinner more nearly first-class in price
than in quality. The narrow hall leading to the dining-room is
crammed with men—Person’s Hotel fed four-hundred a day—waiting
their turn for vacant chairs at the tables. Bolting the meal hurriedly,
the next inquiry is how to get down the creek. There are no coupés,
no prancing steeds, no stages, no carriages for hire. The hoarse
voice of a hackman would be sweeter music than Beethoven’s
“Moonlight Sonata” or Mendelssohn’s “Wedding March.” Horseback-
riding is impracticable and walking seems the only alternative. To
wade and flounder twelve miles—Oil City is that far off—is the dreary
prospect that freezes the blood. Hark! In strident tones a fierce-
looking fellow is shouting: “Packet-boat for Oil City! This way for the
packet-boat! Packet-boat! Packet-boat!” Visions of a pleasant jaunt
in a snug cabin lure you to the landing. The “packet-boat” proves to

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The Evolution Of Exudativory In Primates Anne M Burrows Leanne T Nash

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The Evolution Of Exudativory In Primates Anne M Burrows Leanne T Nash

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