Inia geoffrensis in Captivity in the United States Melba C. Caldwell and David K. Caldwell Marineland of Florida, Route l, Box 122, St. Augustine, Florida 32086, U.S.A

and Randall L. Brill Chicago Zoological Society, Brookfield Zoo, Brookfield, Illinois 60513, U.S.A.

Abstract The history of Inia geofrensls in the United States covers a period ofthree decades which is characterized by the combined acquisition of as many as 70 animals over the first ten years, an abrupt decline in importation, and a loss of specimens leading to the current captive population of one. Data obtained from several institutions, availablo literature, and the personal files ofthe senior authors are reviewed. Such factors as the hazards oftransport, longevity, maintenance conditions, necropsy findings, and com- parisons between Inia and the more familiar marine species, Tursiops truncatus, are considered. Insti- tutional data are additionally summarized in tabular form. The authors suggest that crowding andlor the overly aggressive behavior of Inia kept in groups contributes to the species'poor longevity record in captivity and that isolated individuals or animals that can be readily separated have a greater chance for survival in a captive environment.

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Institutions and attractions that have maintained Inia in the Historical Background United States for public viewing include the Fort Worth Zoolog- In 1956, Silver Springs of Ocala, Florida, sent an expedition ical Park in Fort Worth, Texas; Homosassa Springs in Homosassa to South America which brought back four Amazon River dol- Springs, Florida; the Aquarium of Niagara Falls in Niagara Falls, phins, Inia geffiensis, captured in the Upper Amazon River in New York; the John G. Shedd Aquarium in Chicago, Illinois; the the vicinity of Leticia, Colombia. These four animals became the Milwaukee County Zoo in Milwaukee, Wisconsin; the Pittsburgh first live dolphins of their kind to be imported into the United Zoo in Pittsburgh, Pennsylvania; the Steinhart Aquarium in San States. Although one was dead upon arrival and another lasted Francisco, California; Sea World in San Diego, California; Busch only a day, the remaining two dolphins lived for over a year and Gardens in Tampa, Florida; the Toledo Zoo in Toledo, Ohio; the were observed and monitored by Layne and Caldwell (1964)' Crandon Park Zoological Gardens in Miami, Florida; Marineland Public aquaria and oceanaria apparently found these mammals of Florida in St. Augustine, Florida (Fig. l); and Marineland of ofinterest for, in the late 1950s and early 1960s, several either the Pacific in Los Angeles,r California. sent expeditions to South America or purchased one or more Several animal importers included Inia in their collections as specimens from animal importers. The freshwater habitat of these well and often provided public access to view them. These in- dolphins enabled the aquaria to take advantage of the public cluded the Tarpon Zoo of Tarpon Springs, Florida; the Monte Fish interest in dolphins without having to build new and more costly Vista Zoological Park in Bloomington, California; the Gulf in Vero saltwater facilities. The marine oceanaria could also expand their Hatchery in Palmetto, Ftorida; the Paramount Aquarium Shippers in Miami, Florida' The collections of cetaceans with a new and somewhat unusual species. Beach, Florida; and Animated expeimental studies The public, as it turned out however, never really distinguished United States Navy also maintained lniafot and later on the Inia from the more familiar marine dolphins. Their popularity at their facilities at Point Mugu in California for a short period of time' quickly waned and oceanaria did not continue to maintain them Hawaiian island of Oahu, both Caldwells' records, as of 1965, out of the 34 for very long. In the Caldwells' experience, when Inia were at the According to the been imported into the United States up to that height of their popularity for institutional display more than one- Inia thal had only four were alive (Herald and Dempster,1965). Just one suc,L institution would have continued to bear the high costs of time, the of animals imported had risen to importing and maintaining these unique dolphins had public in- year later in 1966, total 70, of which 19 were alive (Herald, 1967). We have terest supported it. more than 35 f t I

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Figure 2. ,,Golf-ball disease" shown in the center of this photograph of a live adult captiye Inia geofriensis is a common ailment of this species in captivity. Many ofthe small scars are the result ofwounds inflicted through aggressive actions from other captives ofthe same species. (Photograph courtesy of Marineland of Florida.)

the interest in Inia is not likely to increase to the point of en- well be related overall to what has been described as the "golf Figure Originally preilicted to be an advantage for success in captivity, the keeping of such large groups of Inia geofrensis later proved to be a 1. couraging the importation of them in any significant numbers. ball disease" (Klocek, 1981; Pier and Madin, l9l6 as cited by problem (Photograph by William A. Huck, courtesy Marineland of Florida.) because of aggressive behavior. Best and da Silva, this volume) (Fie. 2). Both lung and skin problems encountered with Inia have, in some cases, been cor- Conditions and Longevity in Captivity no figures for the total number of imports as of this writing ( I 986) Transport rected by the use ofantibiotics ofchoice. but can report that only one Inia ctrcently survives at the Pitts- The institutions from which data have been obtained account The 32.6 month average captive longevity that we have cal- burgh Zoo. For whatever the reason, the rate of importation of Before presenting the data on captive Inia, a brief word on for a variety of captive conditions. These range from open-air culated and the necropsy findings listed in Table l, however, may the species rose rapidly and then abruptly declined to none. Leg- transportation is in order. Those first dolphins brought to Silver artificial pools and natural springs to partially and completely not reflect the true state of affairs. We believe that the most serious islation regarding marine mammals and endangered species in Springs, Florida in 1956 had difficulty surviving the long, high- enclosed artificial environments. The maintenance conditions are health problems relative lo Inia held in captivity are stress and the United States, as well as the high costs involved in acquisition altitude flight (Allen and Neill, 1957), as did the animals brought described in Table I where available. injury induced by aggressive behavior between animals (Fig. 3). of these animals, have certainly contributed to this decline since back from Iquitos, Peru by Marineland of Florida in 1966. Herald We have calculated longevities of the animals listed in Table Much of the aggression is of a sexual nature. The files of the senior the early 1970s. (1967) mentioned similar problems as well and Dempster (1965) I. Dolphins that were shipped to other institutions at some point authors are replete with accounts of sexual attacks by males on described the hazards of a comparatively shorter transport from after arrival at their initial destinations were excluded from the females, on young of both sexes, and infants, including attacks Florida to California. Presumably, transport problems related to calculation as were those dolphins that were on one dead infant. We no longer believe that a social breeding Data Base of average longevity, flight, altitude, and duration are characteristic ofthe species and motherless calves, dead upon arrival or dead within 24 hours colony ofthis species in captivity, which appeared to be a desir- The present report is based on the records provided by six we suggest may be alleviated by the administration of oxygen after arrival. We eliminated these latter cases on the grounds that able situation in the mid- to late 1960s, is possible. The attacks cooperating institutions that have held Inia, the literature, and and/or cabin pressurization, keeping the animals warm in flight, death as a result of transport or a failure to wean does not fall are to.o violent, putting some dolphins into shock and leaving on the Caldwells' personal files. As might be expected, over the years and the initiation of a prophylactic antibiotic program prior to within what would normally be expected relative to captive life- others gashes which are prone to infection. It is apparent to us for the data at many institutions, all or in part, have been lost and transport because experience has shown that these dolphins are span. The average captive longevity of the 35 remaining animals that the fewer animals kept together, the better their chances what is available is, in many cases, dimcult to interpret. Because prone to respiratory distress often leading to pneumonia and tend considered was 32.6 months. captive survival. problems aggression. The of these and other limitations, we are restricting this report to to be more subject to infection from cuts and abrasions often Contributing causes ofdeath are listed in Table I and cover a Most aquaria reported relative to 47 copulations associated material obtained from institutions within the United States. suffered in handling than marine dolphins. Methods of handling wide spectrum. In addition to those causes listed, gall stones and Fort Worth Zoological Park reported large male directed toward a female within However, the species has also been held in both Europe and Japan have, of course, improved since those early days when Inia were erysipelas (for a description of this condition see Geraci et a1., with aggression by a twenty-three minutes (McCusker, (Best and da Silva, this volume; Tobayama and Kamiya, this being widely imported, and consequently were the species more 1966) have been reported in captive animals. "Lung problems" a period of three hours and resolved by the placement of a fence volume; Collet, 1984) as well as South America. Even so, the popular today the success in transporting them might be expected are prevalent in wild Inia (Best and da Silva, this volume) and 1913). That situation was Springs similarly separated a following information should be helpful in bringing together what to be greater. However, the public is now more attuned to exciting pneumonia similarly appears to be the disease to most often attack between the two dolphins. Silver means of a fence (Layne and is available. and athletic shows with marine dolphins and killer whales and otherwise healthy captive animals. Reports of skin problems may large male from a smaller male by 36 37 Table I Conditions of Inia Captivity and Necropsy Results at Six Different Locations

Number Water Animals Tank Size in Gallons Treatment Water Air Fish Food Insti- in Data Water and/or (chlorine Tempera- Tempera- Pounds/Animal Necropsy Findings (may include tution Base Dimensions in Feet in ppm) ture in oF ture in'F and Typer several findings) A 2l 44 x 22 x 6"7" plus2 0.4 72-73 ambient 6-12 nephrosis or other kidney problems connecting ofca. 10.5 but en- capelin, herring, (5), "golfball disease" (l), scolio- -F x 6 each, total ca. closed and blue runner sis (2), skin problems (2), pneu- 44,000 plus gallons monia (4), biliary inflammation (l), foreign object ingestion (1), subacute endocarditis (1), nothing found (3); not necropsied (4) B 2 50 x l9 x 5(maxi- not not ambient not known not known, but life spans included mum) known known but out- in averages doors C 5 30x9.75x5(maxi- 0.1 5-0.4 77-84 ambient 6 nephritis, epidermal infection, mum) but en- mackerel, herring spleenitis, peritonitis, bile duct closed hyperplasia, septicemia (3), bac- terial pneumonia D 2 initiallyca. 15 x 15 x 0.7 67-72 74-80 6-7 recurrent skin problems, "Golf ball 6 with 6000 gallons, butterfish disease," teeth wom to gums later 13,000 gallons E 3 20x30x6(maxi- not 80 76-80 7-8 occluded intestines, ulcers, congest- mum) known smelt, Spanish ed liver, septicemia, thickened mackerel, her- bladder wall, congestive heart ring failure F 2 900x20-25x4-5 none 72 ambient 10-17 pneumonia (2), brain hemorrhage, (maximum) (preferred blue runner Heterotrema caballeroi found in deeper pools). Later lungs and trematode eggs in brain moved to pen 75 x of one animal 40 x 5'10"

' Food fishes: blue runner (Caranx crysos), butterfish (Peprilus tiacanthus and P. burtf, capelin (Mallotus vll/osas), herring (Clupea harengus, Opisthonema oglinum), mackerel (Scomber scombras), smelt (Osmerus mordax), Spanish mackerel (Scomberomorus maculatus). Many institutions administer dietary supplements, usually multivitamins, in capsules or pills inserted into some of the whole food fish.

Caldwell, 1964). The Toledo Zoo reported that a male became after another small male was introduced into its tank resulting in belligerent toward a smaller female who survived for only three a desirable weight gain for both dolphins. Although more positive, weeks (Max Hofmeister, pers. comm., 1965). At Sea World in these instances seem to be the exceptions rather than the rule San Diego, a large resident male nearly knocked a newly relative intro- to the keeping ofgroups of Inia in captivity. Best and da Figure 3. A small fiemale Inia geofrensh, newly introduced into a tank at Sea World in San Diego, California, is almost knocked out of the water duced female completely out of the water (Caldwell and Caldwell, Silva (this volume) do report that, although most frequently ob- by a larger, aggressive resident male. (Photograph by David K. Caldwell') t961). served to be a solitary feeder, Inia may join in apparently loosely The three Iniathat have thus far survived the longest in cap- organized groups to cooperatively herd food fish in the wild, but tivity were the last survivors of the groups in which they were such cooperative behavior has not been observed in captives. phone (Caldwell et al., 1966). At any rate, Iniahabituate to novel delivered or kept and have been maintained as individuals re- While behavioral problems due to aggression have been reported Behavior Miscellaneous objects and can be trained to perform relatively simple behaviors spectively at the Aquarium of Niagara Falls, for almost 20 years; for the marine Atlantic bottlenosed dolphin, Tursiops truncatus geffiensls for the public, as was the case at the Fort Worth Zoological Park. the Shedd Aquarium, for l6 years; and the Pittsburgh Zoo, for (Caldwell et al., 1968; Caldwell and Caldwell , 1977), we do not In comparison to marine dolphins, Inia is a fairly Such behaviors were, however, limited by the physical abilities 19 years and still alive at the time of this writing (1986). Another believe that they usually reach the degree or intensity of those lethargic species. It usually swims rather leisurely in its tanks as single animal lived over l7 years in Japan (Tobayama and Ka- seen irr Inia. opposed to the high levels of activity commonly associated with of the species. 'fursiops On the other hand, sexual activity in Inia compares more closely miya, this volume). The longevities of these individuals so greatly Correlations between longevity and group sizes still require truncatus. This characteristic is reflected in its rate of with its marine counterparts. Such behavior includes a great deal exceed those of the other captive Inia thal we find it difficult, proper analysis but the general tendency is reflected in the fol- food intake as well. Atlantic bottlenosed dolphins are known to homosexual and heterosexual activity indeed, to merely attribute their success to the "survival of the lowing dala lrom one institution: eonsume about 15 pounds offish per day compared to the 6-8 of masturbation and both quite active and lengthy in medically fittest." We feel that there has been a significant number pounds of fish per day consumedby Inia (Table 1). (Spotte, 1967), all of which can be of Inia in captivity which of other Inia kept in groups with excellent medical care and After a short period of acclimation, captive Inia have laken duration. There have been three births Animals received in one shipment numbered five. that lived only moments at the husbandry whose lifespans do not begin to approach those ofthe food from human hands quite readily. At least one institution include one stillbirth and another Animals dying in transport numbered one. 1973) three animals cited above. took advantage ofthe situation and provided food fish for a price Fort Worth Zoologtcal Park (Huffman, 19701' McCusker, Animals dying within the first twenty-four hours numbered Marineland of Florida in which the infant In addition to the incidence ofovert aggression among Inia, a to tourists who were then allowed to feed the animals. Food for and one live birth at one. weeks (Caldwell and Caldwell, 1972). large male at Marineland of Florida was reported to steal, but not marine dolphins has been provided in this manner in several survived for two have demonstrated the frightening habit of eat, food from a smaller male even though the smaller male tried Of the three remaining animals: institutions based on the grounds that both tourists and animals Some captive Inia (sleeping?) upside down at the bottoms of their pools to hide its fish. Obviously, sick individuals are placed at an even enjoy the interactions and that the costs offood, as well as research resting programs manner. (Caldwell eta1., 1966; Herald, 1969). They were so immobile that greater disadvantage when exposed to aggression or prevented The first to die lived twelve months leaving two in the tank. in some cases, can be alleviated in this placed their tanks they appeared to be dead. Observers that were unfamiliar with from obtaining food. On the other hand, Johnson (l 982) reported The second to die lived 32 months, leaving an isolated Inia will apparently play with objects in survi- this kind of behavior often frantically notifled the institution's food sharing behavior by a male Inia toward a female who re- vor. (Herald and Dempster, 1965; Klocek, 1981). Although Herald reactions of fear to novel staffwhenever it was seen. Pilleri (1969) reported a similar death- jected the food at the Pittsburgh Zoo. This same large male was After 41 months, the surviving dolphin was shipped to another and Dempster (1965) report initial have reported fearlessness in like appearance for captive Iniathat rested at the bottom on their also reported to have fed a smaller male. Herald (1967) reported institution while healthy and survived for an unknown length objects pliced in the tank, others bellies. that a small male at the Steinhart Aquarium ate more heartily of time. response to such events as the introduction of a strange hydro- 39 38 individuals in the wild and most sightings are of solitary animals Inia geffiensis. Int. J. Syst. Bact. 26(4),93-99 (as cited by Best and Woodard, J. C., S. G. Zam, D. K. Caldwell and M. C. Caldwell. 1969. Research volume). Some parasitic diseases ofdolphins. Path. Vet. 6,257-272. (Best and da Silva, this volume), the high rates of aggression in da Silva, this Pilleri, G. 1969. On the behavior of the Amazon dolphin, Inia geof- Zam, S. G., D. K. Caldwell and M. C. Caldwell. 1970. Some internal Most of the research that has been conducted on Inia in lhe captivity possibly resulted from keeping animals in enclosures frensis, in Beni (Bolivia). Rev. Suis. Zool. 76, 57-91. parasites from freshwater cetaceans from the Upper Amazon River. United States has focussed on their sound production and hearing too small to allow them to maintain normal spacing between Spotte, S. H. 196T.Intergeneric behavior between captive Amazon River Invest. Cetacea 2, 250-251. et al. (1966) no more than two individuals be capabilities. Their sounds were discussed by Caldwell individuals. We suggest that dolphins Inia and Sotalia. Underw. Nat. 4(2),9-13. and Caldwell and Caldwell (1970). The anatomy of the eye has kept in the same enclosure. Inasmuch as experience has suggested been examined by Dawson ( I 980) who compared Inia wilh other that it is very possible that antagonistic behavior may occur be- cetaceans. Harrison and Brownell (1971) studied the gonads of tween two or more individuals, a fast and effective means of 18 of these captive dolphins, and Zam et al. (1970) reported separating them should be provided. internal parasitic trematodes and nematodes. Woodard et al. (1 969) earlier reported pulmonary trematodiasis. Data based on behav- Literature Cited ioral observations have been contributed largely by the curatorial staffs of the various institutions in which Iniahas been held and Allen, R. and W. T. Neill' 1957. White whales of the Amazon' Nal' are invaluable. These same people have been instrumental in Hist. 66,324-329. in the Amazon dolphin, initiating investigations into disease related problems' Importers, Caldwell, D. K. 1966. Morphological variation geofrensis (Blainville). Yearbook of the American Philosophical aquaria, and oceanaria were very cooperative in making the car- Inia Society 1966, 328-331. caises of animals that died in their care available for scientific Caldwell, M. C. and D. K. Caldwell. 1967. Intraspecific transfer of Skeletal materials, especially skulls, thus found their way study. information via the pulsed sound in captive odontocete cetaceans. In: the U.S. National Museum of Natural into museums including Les Systemes Sonars Animaux, Biologie et Bionique,2' R'-G' Busnel Museum History in Washington, D'C.; the Los Angeles County (Ed.), p. 879-936. Laboratoire de Physiologie Acoustique, Jouy-en- Florida State of Natural History in Los Angeles, California; the Josas, France. Museum in Gainesville, Florida; and the California Academy of Caldwell, M. C. and D. K. Caldwell. I97O- Further studies on audible geffiensis' Los Sciences in San Francisco, California. Skeletal materials are also vocalizations of the Amazon freshwater dolphin, Inia in the collections of the American Museum of Natural History Angeles Co. Mus Contr. Sci. 187' 1-15. 19'72. The littlest uglv dolphin' in New York and the Field Museum of Natural History in Chi- Caldwell, M. C. and D. K. Caldwell' Frontiers l8(l), 24-29. cago, Illinois. Systematic studies were initiated during the period Sea Caldwell, M. C. and D. K. Caldwell. 1977. Social interactions and in wfrlcir so many individuals of the species were being imported reproduction in the Atlantic bottlenosed dolphin' ln: Breeding Dol- States (Caldwell, 1966). into the United piins: Present Status, Suggestionsfor the Future' S' H' Ridgway and k. Benirschke (Eds.), p. 133-142. Marine Mammal Commission Re- PB- Summary port No. MMC-76/07, National Technical Information Service, 273 673. A brief review of the history of the Amazon River dolphin, Caldwell, M. C., D. K. Caldwell and W. E. Evans' 1966' Sounds and Inia geffiensls, in captivity in the United States indicates that behavior ofcaptive Amazon freshwater dolph\ns, Inia geoffrensis' Los similar husbandry problems were encountered in at least six dif- Angeles Co. Mus' Contr. Sci. 108, l-24' and B. C. Townsend, Jr' 1968' Social ferent facilities. The average longevity in captivity for the animals Catowitt, M. C., D. K. Caldwell considered was 32.6 months and the major causes of mortality behaviorasahusbandryfactorincaptiveodontocetecelacearns,Proc. Second Symposium on Diseases and Husbandry of Aquatic Mammals' were: Marineland Research Laboratory, St. Augustine, Florida, p' 1-9' l. Transportation (shock and pneumonia). collet, A. 1984. Live captufe ofcetaceans for European institutions. 2. Stress and injury brought on as a result of aggressive be- Rep. Int. Whal. Comm.34,603-60'7. eye. In: Cetacean Behavior: Mech- havior between individuals (see discussion below)' Dawson, W. W. 1980. The cetacean and Functions- L. M. Herman (Ed'), p' 53-100' John Wiley 3. Skin problems, many arising from the bites and blows in- anisms and Sons, New York. flicted by other individuals or by the rigors oftransport' Dempster, R. P. 1965. Shipping a live Amazon dolphin' All-pets 36(3)' unusually high percentage of kidney 4. Pneumonia and an 13-15,17. to that found in Tursiops truncatus' problems as compared Geraci, J. R., R. M. Sauer and W' Medway. 1966' Erysipelas in dolphins' Res.27(ll7)' 597-606. The captive behavior of the animals at the six institutions Am. J. Vet. Harrison, R. J. and R. L. Brownell, Jt. 1971. The gonads of the South providing data for this review, as well as animals not considered American dolphins, Inia geoffrensis, Pontoporia blainvillei, and So- Table l, indicates a high level oflethargy unlike the captive in talia fluviatilis' J. Mammal' 52(2), 413419' species. While it is apparent behavior of the more familiar marine Herald, i. S. tSOl. Boutu and Tookashee-Amazon dolphins' Pac' Dis' thal Iniaare trainable, they cannot accomplish the more complex 20(t),2-e. and acrobatic behaviors commonly seen in such marine species Herald, E. S. 1969. Aquatic mammals at Steinhart Aquarium' Pac' Dis' as Tursiops truncatus. 22(6),26-30. Iniakiptin groups may be very aggressive toward each other, Herald, E. S. and R. P. Dempster. 1965' Meet "Whiskers"' Aquar' J' particularly with respect to sexual behavior. In the Caldwells' 36(s), 2 | 3-2t 5, 23 6-23 7 . W. 1970. Notes on the first captive conception and live experience sexual harassment by the largest males constituted a Hutrman, E. dolphin in North America' Underw' Nat' 6(3),9- major problem in a social grouping of these dolphins' Sexual birth of an Amazon 11. biting by a large, sexually mature male was a threat to even a Johnson, R. H. 1982. Food-sharing behavior in captive Amazon River and she had to be separated to prevent large, mature female, dolphins (Inia geffiensis). Cetology 43, l-2' mature male also was guilty ex6nsive damage by bites. This same Klocek, R. 1981. Chico's story: a special dolphin. Aquaticus l3(2), 1-9' behavior toward a juvenile male, cutting ofthe Amazon dolphin, of intensive aggressive Layne,'Inia J. N. and D. K. Caldwell' 1964' Behavior the latter badly. An immature male was aggressive in biting other geofrensLs (Blainville), in captivity. Zoologica 49, 81-108, pls' animals and preventing a juvenile female from eating. Both male I-IV. and female Iniu artacked a recently-introduced small juvenile McCusker, J. S. 1973. Observations on the Amazon Dolphin, Inia geof- (Unpubl' ms') male, and a sick female was the subject of intensive sexual har- frensis, in Captitity at the Fort Worth Zoological Park' assment by four males of assorted sizes until she could be sepa- Pier, C. B. and S. H. Madin. 1976' Streptococcus iniae, sp. nov', a beta- rated. As this dolphin rarely occurs in groups of more than two hemolytic S/rep tococcus isolated from an Amazon freshwater dolphin' 40 41