Journal of Abnormal Primatology (1990), 29(2):31-44. |
Comparative Plummeting Rates in Ten Species
|
Abstract. Extensive field data has now been collected on the plummeting rates of ten species of Old and New World primates. Approximately 6000 observation hours was devoted to each species, five from South and Central America, and five from Africa and Asia. Emphasis was placed on jump-off posture and plummeting mode, and data recorded included height and duration of plummet, time to reach terminal velocity, and speed at terminal velocity. Observations were made by the authors and by a skilled team of trained natives who were accustomed to watching things. The landing was not part of the study, but there were 87 observed fatalities and 526 other injuries. In general, the larger primates took a longer time to reach a higher terminal velocity, but on the whole displayed a greater variety of adaptations designed to affect their plummeting rates and lessen the stress of impact. Infants of all species were rarely capable of achieving the plummeting rates of adults. |
As long
as men have watched primates and primates have peered obtusely back, there
have been many unasked questions in the murky field of behavioral science,
and one of the most persistent has always been "Which monkey species
plummets the fastest?" Ever since Plato's Why Falls the
Simian?, curious minds have wondered whether the lesser primates actually
do plummet more slowly than the great apes, and whether this is due to
lesser weight, greater surface area-to-volume ratio, both, or something
else entirely. Plato spent several years pursuing primate studies,
and was a great fan of monkeys in general. It has never been clear
whether he actually ever saw an ape, but it seems certain that either
a giant pygmy chimpanzee or a gorilla of some sort or other was the animal
he referred to as Primator impericus, the 'royal monkey.' |
It is
not always clear just what ambition any given primate is attempting to
realize by plummeting in this fashion, what inhuman emotion drives him
to risk his life and limb this way, or what warped evolutionary pressure
created this odd behavioral pattern in the first place, but there seems
little question that it is widespread across the primate order, and possibly
present also in species that are now extinct, such as Robertson's pink-nosed
guenon and the Alpine wheezing macaque (Shaw, 1979; Axelrod-Abernathy,
1982; Winklehurter, 1990). While it is true that there may be specific
non-evolutionary reasons for plummeting, such as escaping predation by
upper canopy hunters, trying to beat an opponent to a terrestrial food
source, or simply losing one's balance and plummeting inadvertantly, the
ubiquitous nature of primate plummeting in most forest environments in
both the Old and New Worlds demonstrates that it is a behavioral pattern
that has been around for a long time. |
Species we were able to research thoroughly. |
New World: |
Ateles olympica(jumping spider monkey) Cajuru imperator (purple wannaby) Procebus croesusii (Croesus monkey) Andradus blanca (white-cheeked muscatel) Saguinus rex (blue-blooded tamarin) |
Old World: | Cercopithecus subterraneus
(bluetail guenon) Nasalis nasalis (hairy-nosed proboscis monkey) Nycticebus slowpokeii (really-slow-loris) Tarsius irritatus (sulky tarsier) Presbytis oro (gold leaf monkey) |
Once the species choices had been made, we still had to arrange hotel reservations and flight plans, update our passports, cancel the mail and turn off the electricity, notify our next of kin, and get all our shots. We each had to close down the work we were doing at our respective institutions (for me it was the spitting behavior of bluetail guenons, and for Dr. Knuckleberry it was camouflage adaptions of the Alaska snow mouse), all the while faxing itineraries back and forth like frisbees. Once in the field it was not always easy to collect the kind of data we wanted. The study began somewhat intermittently and then months of luckless searching would sometimes go by before we would suddenly see a flurry of plummeting activity, and limping primates could be seen from morning till night. Except for the Callitrichidae, most primates apparently preferred not to plummet on weekends, so this left us free to enjoy ourselves without fear of losing valuable observation time, and by Friday afternoon we were usually all too willing to pile into whatever kind of transportation we could get and head for the nearest place that sold hot Madagascar toddies or oilberry beer. We also established to our satisfaction that most primates rarely plummet on purpose in bad weather (although bad weather does increase the rate of inadvertent plummeting), so whenever the rainy season started in one place, we would immediately move on to another. This kept us on the go quite a bit, but the result was a lot of really terrific data on primate plummeting rates, and measurably lower laundry bills. Even in good weather, plummeting is an on-again-off-again type of behavior. Besides which, there are many specific and sub-specific differences in the rates at which females engage in plummeting as opposed to males, infants as opposed to adults, and residents as opposed to migrants. Primates are often hesitant to jump when they know they are being observed, and occasionally will refuse to plummet altogether. My co-author once actually had to climb a tree to push a reluctant primate out of its nest so that we could time its descent and finish off a block of data, but thankfully that was not something that we had to resort to very often (Knuckleberry, 1989). Of the plummet as a whole, it was the jump-off that was most difficult to observe, since it usually took place high in the upper canopy, and better observations are usually only made when the animal in question comes crashing through the mid-story about 10-15 meters from the ground. On certain occasions, the first warning that the observer has of a plummeting primate is the characteristic shrill screech that many primates make as they approach the ground. While there are species that have been seen jumping from the lower branches, it is perhaps not surprising that primates as a rule typically seek to maximize their plummeting efforts by reaching terminal velocity as early in the descent as possible. What little information is known of pre-plummet postures and the actual jump-off itself has been assiduously gathered by natives who are able to climb high enough to observe it. |
Before
we can discuss actual plummeting rates and the reasons why those rates
vary so widely, we need to look at the jump-off, because the jump-off
is usually the most crucial phase of the plummet. I have defined
the jump-off in my textbook Primate Pesematology as the first 1.5
seconds of the plummet. It is during the jump-off that the primate
customarily chooses its plummeting orientation, thereby setting the stage
for the plummet to follow. Most primates lack the felid's easy ability
to change its orientation during a fall, thus they are quite keen on getting
the right jump-off angle at the start, and each primate species that engages
in this behavior has its own characteristic pre-plummet stance. |
Fig. 1. Sub-adult male Cajuru imperator
in cloud forest on the slopes
of volcanic Mt. Tehuatán, about to plummet from the upper branches of Acambaro variegata. |
type of plummet utilized by the purple wannaby. Compare this to the very different pre-plummet posture of Procebus croesusii (Fig. 2), with its head thrust defiantly forward, its arms extended outward and upward, and its feet positioned uncomfortably close together. Tarsiers prefer to jump off quietly and unobtrusively, almost sadly, from a vertical clinging position, while Andradus stands quadrupedally on a branch and rotates flamboyantly downward with its tail in the air as the plummet begins, letting go first with the hands, and then with the feet. Gold leaf monkeys have been observed standing bipedally and jumping off to plummet downward feet first, and they also demonstrated a marked preference for plummeting from the crowns of the giant emergent trees Landacia iticulata, Caudatia spoelii, and Provergatus lacchus. The really-slow-loris is slow and deliberate, backing and filling on one branch after another, stopping for long moments to gaze around and then moving again, and generally delaying the jump-off as long as possible. This is especially true during periods when the moon is full. It is not |
Fig. 2. Alpha male Croesus monkey with
head thrust defiantly forward
making a last check of its surroundings before making an early-morning plummet from Ariavaria immensis. |
unusual that a plummet of five seconds is often preceded by hours of pre-jump-off preening and posturing. Ateles is the only primate studied thus far that plummets from a tail-hanging position, while the hairy-nosed proboscis climbs out on the thinnest branch tips before lunging precipitously just as the branch breaks under its weight. Unlike other tamarins, Saguinus rex has evolved the vine-swinging plummet, in which the animal uses a liana to swing back and forth, thus picking up speed and enabling it to reach terminal velocity more quickly. A rare sub-species, S. rex domesticus, has become adapted to living on the roofs of buildings, and has been seen making plummets of up to ten stories in height. Unfortunately, city pavement is less forgiving than forest soil, and after several years of difficult investigative work, our assistants have told us that many of the veterans of these higher suburban plummets are no longer around. |
For the
purposes of the immediate study, the plummet shall be defined as the period
between the jump-off and the landing. To begin with, the typical
primate plummets in one of only two possible modes. He or she either
strikes or is struck by intervening vegetation, or falls straight down
unimpeded. It should be obvious then even to pre-seventh graders
that the bioarchitechtural character of the forest has much to do with
plummeting rates. Further, different workers in the field have interpreted
plummeting rates very differently, choosing for instance to emphasize
duration of fall rather than velocity. Marshall et. al.'s
classic monograph on vegetational morphologies and plummeting rates in
the first issue of the Darwin Society Journal established clearly
that hitting branches and other biotic impediments both slowed the speed
of the fall and increased its duration. Striking intervening vegetative
strata also has possible medical consequences. |
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|
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with its eyes closed like a crazed
skydiver. The gold leaf monkey, on the other hand, ties itself to
a network of green thread vines which sometimes acts to mitigate its headlong
plunge into the dim and uninviting waters of Bali-Bali's countless shrimp
lakes. The prosimians represented in our investigation, Tarsius irritatus and Saguinus rex, are of such diminutive size and weight and have such a high surface area-to-volume ratio to begin with, that they accelerate slowly to reach a moderately sedate terminal velocity, and they both seem content to accept their lot as the slowest of nature's plummeters studied thus far. Nycticebus slowpokeii steps uncertainly off its branch and falls hind-end first with a glazed expression on its face and its arms clasped tightly about its abdomen. |
The landing The landing is defined as the final second of the plummet. It is really not all that important to the subject of primate plummeting rates because, after all, at the point of landing the plummet is over, and at that point plummeting rates of all species tend be zero. The only exception to this is the great blue marmoset, an animal for which data was not sufficient to include in this study, which often chooses to land on a steep slope, prolonging the plummet as it rolls toe over toothcomb down through typical Amazonian hillside vegetation, and giving it by far the longest plummeting duration times recorded to date by western scientists. |
The analysis Having described in some detail several of the gross behavioral features of our sample of primate plummeters, we can now turn to a statistical analysis of the data so painfully gathered over the course of the last two hundred months. Appendix E shows rates of acceleration and terminal velocities for the ten species in our study. It seems clear to us at least that there is no apparent significant correlation between weights, body sizes, and plummeting rates, but of course we could be wrong. |
Conclusions After nine years of laborious planning and eight years of burdensome field work, we have determined almost categorically and without very much doubt that primates do not plummet at exactly the same rate. We tried to use quantum point analysis, and made a brief stab at differential time-distance equations, followed by a bit of isopropic calculus and game theorums (Shaw, 1982), and we utilized to somewhat good effect Professor Ashram Suleiman's fractional data scattering codes as proposed for jumping rates of Turkish viverrids. Our conjectures may be questioned by some, but no one can deny our heartfelt and unfeigned approach. We expect that there may be some negative reactions in the primatological journals, but we plan to press ahead and not allow the naysayers to deter us from our final goal, the publication of which we are planning to be coincident with the 200th anniversary of the War of 1812. |
Acknowledgements The authors wish to gratefully acknowledge the significant contributions, unstinting support, and just darn good cheer and unquenchable high spirits of the following individuals and institutions: Mrs. J. Muffet Gloverhouse, the Icelandic Primate Rehabilitation Project, everyone at the Mahatutu Injured Leaf Monkey Shelter, Dr. Heckelen Jeckel, the Tremblay-Waxenhauser National Center for Primate Disorders, the Russian national primatology newspaper Primatsiya Primatsiya, the Alaska School of Orthopedic Veterinary Medicine, the Antigua and Barbuda College of Caribbean Primatology, Dr. Jerry Archbibble and the Hellmouth Municipal Zoo and Exotic Animal Crematorium, the Natural Geographic Society, the Cheesequake Municipal Man and Mammal Museum, the National Primate Bowel Clinic, Dr. Oondóué M. Boué, Dr. Watanabe Kibombo, Dr. Ambato Ambilobe, and Dr. Miedzyrzecz von Czechowice-Dziedzice. We especially want to recognize the work of the Free Fall Club of Jujube and Togobogo, the French Monkey Academy Precision Flag Plummeting Team, Senhor Teófilo Afonso Rosario Sobradinho, Piet Mons Apeldoorn, the Rochefort-Chateauroux Institute for Simian Science, Drs. P. Rudyard Kiplinger and Buffy Rockefeller, the Hellmouth Human Diseases and Primate Testing Facility, Sir Ian Spotswood Allenby Crofford-Wiggles, the Bluetail Foundation and the Udon Sawan Lesser Ape Sanctuary. We also wish to recognize the contributions to our work of the International Plummeting Recovery Center and the Arizona Science Foundation. We are forever in the debt of Dr. Francois Quimper Bonnetable Rochefort-Chateauroux, Dr. Mawbanna Waddamana, Dr. Oscar Simon Bolivar Bolivar-Fuentes, Mr. George Jefferson, and Dr. Poon Sandandtundra. Finally, we must thank Professor Rolf Sigurd Vanhammerfest, the Baseball Commission, Sigsbee Junior Night College, Primate Nooz and PRIMATE LIFE, the Society for the Prevention of Falling Mammals, the Swedish Primatological Association, the Antique Sulky Tarsier League, Professor Mitsuo Ohhohoho (wherever he is), the people of Afghanistan, and Mr. Christopher Shaw. |
Appendix A Comparative weights of selected species: |
Species |
No.##
|
Grams
|
No.##
|
GrGramsams
|
Ateles olympica Cajuru imperator Procebus croesusii Andradus blanca Saguinus rex C. subterraneus Nasalis nasalis Nycticebus slowpokeii Tarsius irritatus Presbytis oro |
31 19 12 27 8 14 3 21 11 9 |
5,620-6,140 2,775-3,410 6,905-8,195 975-1150(?) 355-585 3,370-5,425 9,275-10,350 1,120-1,190 95-155 4,500-5,865 |
20 12 14 22 13 14 9 13 14 8 |
4,295-5,610 1,435-2,155 4,235-5,360 820-935 285-320 2,725-3,385 6,450-7,230 795-1,055 90-135 3,855-4,120 |
Appendix B Comparative dimensions of selected species: |
Species | No.## | Head/Body Length (in mm) |
No.## | Tail Length (in mm) |
Ateles olympica Cajuru imperator Procebus croesusii Andradus blanca Saguinus rex C. subterraneus Nasalis nasalis Nycticebus slowpokeii Tarsius irritatus Presbytis oro |
12 6 15 21 17 13 4 10 22 9 |
390-545 317-385 436-512 292-367 165-305 392-618 520-723 263-383 93-159 422-780 |
14 10 12 8 24 18 7 13 28 19 |
622-784 426-513 297-340 352-456 325-420 470-981 595-610 --------- 129-273 491-920 |
Appendix C Locations where each species was studied: |
Ateles olympica............................. Cajuru imperator............................ Procebus croesusii........................ Andradus blanca........................... Saguinus rex................................... Cercopithecus subterraneus........ Nasalis nasalis............................... Nycticebus slowpokeii................. Tarsius irritatus.............................. Presbytis oro................. ................ |
Urubupunga Research
Station, Brazil Santa Rubia Island Purple Wannaby Refuge, Gorgonzola Urubupunga Research Station, Brazil Gurupi-Matapao Higher Mammal Preserve, Peru Quiriquiri National Primate Park, Venezuela Makokou Study Area, Gabon Kualakurun Primate Reserve, Borneo Udon Sawan Lesser Ape Sanctuary, Thailand Atapao Monkey Park, Bali-Bali Mahatutu Injured Leaf Monkey Shelter, Malaysia |
Appendix D Coefficients of drag (M and L) and mean frequency parameters |
Species |
M
|
L
|
P1
|
X2
|
Ateles olympica Cajuru imperator Procebus croesusii Andradus blanca Saguinus rex Cercopithecus subterraneus Nasalis nasalis Nycticebus slowpokeii Tarsius irritatus Presbytis oro |
0.032 0.016 0.029 0.007 0.044 0.058 0.018 0.025 0.009 0.030 |
1.68 2.72 3.40 2.97 4.20 2.68 0.99 5.21 4.39 2.22 |
121.5 124.3 98.2 159.2 140.0 101.6 117.8 109.1 125.4 119.7 |
2.2+1 1.9+1 4.0+1 3.9+1 1.2( ) 2.3-1 2.8-1 1.5-2 3.6-2 2.5-3 |
Appendix E Mean weights (MW) and terminal velocities (TV) of selected species: |
Species | MW: (in gms) |
Speed at TV: (in feet/sec) |
Reach TV at: (in feet from jump- off point) |
|
|
|
|
Tangential acceleration is a function of locomotor mechanics, and all figures are means. |
* For comparative purposes |
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