J. Anim, Physiol. a. Anim, Nute, 87 (2003), 66-74 Receipt of is. 03. 06. 2002 @ 2003 Blackwell Verlag, Berlin Accepted: 23. 10, 2002 ISSN 0931-2439 Institute of Nutrition, University of Veterinary Medicine, Vienna, Austria Influence of environmental humidity and dietary protein on pyramidal growth of carapaces in African spurred tortoises (Geochelone sulcata) By C. $. WarsneR and C. Tew Summary ‘The carapaces of captive-raised tortoises (terrestrial chelonians of the zoological family Testudinidge, often develop pyramidal shaped osseous growth centrally within the horny places. With very few exceptions (eg. Geochelone elegans, Psemmobates sp.), this conical wth pattern is considered to be pathologic. This very common defect is believed to be an important indicator of the quality of captive tortoise management. This study was Aesigned to examine the effect of dietary protein level and environmental humidity on the degree of pyramidal growth in che carapaces. Fifty recently hatched African spurred tortoises (G. suleata) were raised for 5 months under artificial conditions of varying environmental humidity and dietary protein content (14% vs. 19% vs. 30% crude protein jn dry matter). Humps of the carapaces that developed and blood values of calcium, phosphorus and haematocrit were measurod and compared among. groups. Dry environmental conditions (24.3-57.8% and 306-74.8% relative humidity) produced taller humps. than humid conditions (45-99% relative humidity). Hump formation differed significantly (p< 0.091) between these three groups kept under different humidity conditions, Variable dietary protein had minor, positive impact on this pathological formation of humps (pyramidal growth syndrome, PGS). Analysis of blood (caleium, phosphorus and haematocrit) offered no further explanation as to the evelopment of the humps. ‘Zusammenfassung Bei in Gefangenschaft gehaltenen Schildkrdien entwickelt sich hiufig im Zentrum der Hornplatten des Ruckenpanzers ein pyramidisch geformtss Knochenwachstum. Mit wenigen Aussahmen (7-B. Geochelone elegans, Psemmobares sp.) wind dieses kegelforige ‘Wachstum als pathologisch angesehen und es besteht die Annahme, dass dieses Wachs- tumsverhalten ein Parameter zur Beurtellung der Haltungsbedingunigen der Tiere ist. In vorliegencem Versuch sollte der Einflu8 der selativen Luftfeuchtigkeit sowie des Rohproveingehaltes im Fuster auf die Hackerbildung uncersacht werden. Punizig Schlupfiage von afrikanischen Sporeaschildkriten (G. slate) warden fat Monate lang, unter snterschiedlich feuchten Umweltbedingungen gehalten sowie mit Furcermitla, welche sich durch ihren EiweiSgehale umterschieden (14% vs, 19% vs. 30% Rohprotein in der Trockensubstanz), emihre. Die wihrend dieser Zeit gebildeten Panverhocker wurden anschlieBend vermessen und die Kalzium, Phosphor- und Himatokritwerte im Blut lntersucht und zsvischen den Versuchsgruppen verglichen, Trackene Haleung (24.3-37.5% Dew. 39.6-74.8% relative Luftfeuchtigreit) ihre ra stirkerer Hickerbildung als feuchte Haltung (45-99% relative Luftfeuchtighet). Die Hockerbildung war zwischen den deci U.S. Copyright Clsrnoe Caner Ce Stein: 931-203972ECNTOI-C04$ 1599/0 www Machwelley acy nflonce of Gruppen, die bei untersch hiedlich (p < 0,001). Der aicht abgesicherren’ Ein Panzerhbckerbildung (Py einen klivenden Beitrag 2 The crapaes of captive Tessin) often devel plates (ig 1) With very Ennial growth patria esprit. Ths deh Indiator of the quay o cxplsin ss phesomeson (0972) sewed thn ch fangs ops eh Sslevm : phosphorus (Ca nave been ugg tel by Made 2200, Wicker, 2009), development of pyramid! fuchar onsodystrophia fo Gisese (BD) Some a pyramidal growth synod {iekson amt Cooper, 14 Tggeescer, 2008 Material may be protected by copyright law (Title 17, U.S. Code)Receipt of Mss 93. 06, 2002 ‘Accepted: 23. 10, 2002 edicine, Vienna, Austria ad dietary protein on san spurred tortoises ) onians of the zoological family owth cenctally within the borny Psamnmaobaies sp, this conical rminon detest is believed 10 be e management. This study was ind environmental humidity on cently hatched Afican spurred ifiial conditions of varying v8. 19% ¥s. 30% crude protein | and blood values of ealeium, smpared among. groups. Dry relative. humidity) process humidity), Hump formation groups kept under different ninor, positive impact on this Srome, PGS). Analysis of blood urther explanation as wo the se sick baufig im Zentrum der mies Knocheawachstum. Mit tex sp.) wird dieses kegelfocmige s Anmahie, dass dieses Wachs- gsbedingungen der Tiere is. In an Luuffeuchrigheit sowie des ; untersucht werden, Finfzig Teata) wurden Fant Monate lang, salten sowie mit Futtermiceln, "s. 19% vs. 30% Rohproveia in bildeten Panzerhdcker wurden und Himatokritwere im Blut ‘Trockene Hlaltung (24.3-57.8% kerer Hiickerbildung als feuehte bildung war rwischen den deei O46$ 5000 www Hacwlidelgreay Ieflence of bumidiny and dive o» pyramidal erowth of corapases o Gruppen, die bei unterschiedlicher Luftfeuchtigheit yehalten wurden, signifikant unterse: Ficdlich {p < 0,091), Der Proveingehale des Furters hatte einen geringen und statistisch nicht abgesicherten Einflu8 auf den Grad dieser pathologischen_pysamidenformigen Panzerhockerbildung (Pyramidal Growth Syndrome, PGS). Die Blutwerteanalyse konnte keinen klarenden Beitrag 2u diesen Ergebnissea liefern Introduction ‘The carapaces of captive-raised tortoises (terrestrial chelonians of the zoolgical family ‘Testudinidse) often develop pyramidal-shaped osseous yrowth centrally within the horny plates (Fig, 1), With very lew exceptions (e.g. Geochelone elegans, Psarmmobates sp.) this conical growth pattern is considered pathological and occurs mostly in tortoises in captivity. This undesirable, but very common. defect is believed w be an importent indicator of the quality of captive corcnise management. The long history of attempts to explain this phenomenon alresdy demonstrates this. As early as 1972, Obst and Meusel (1972) suggested that the humps were the result of a previous rickety condition. A wide range of possible causes have been mcnrianed in the literature, like nutritional imbalances such as excess dietary protein, deficiency of calcium and vitamin D or an improper calcium : phosphorus (Ca: P).'Also rapid growth rate and deficiencies of ultraviolet radiation have been suggested as the primary cause (Frye, 1991; Hightield, 1996; McArthur, 1996 (cited by Mader, 1996); Beyon et al 1997; Kiesche, 1997; Walls, 1997; Sassenburg, 2000, Wiechere, 2000). Most of these authors have not made an attempt to associate the development of pyramidal growth with other very common osseous discases of chelonians such as ostcodystrophia fbros, cickets, secondary hyperparathyroidism or metabolic bone disease (MBD). Some authors, however, have either exp icitly or implicitly suggested that pyramidal growth syndrome (PGS) is, in fact, due to the same cause as these diseases ackson and Cooper, 1981; Gabrisch and Zwart, 1998; Kohler, 1996; Kirsche, 1997; Figgenschwsler, 2000; Sassenburg, 2009; Wicchert, 2080). moze thorough understanding Fig. £ Allabra giant tortoise (Genehefone dessamier) with exemsive hump development Material may be protected by copyright law (Title 17, U.S. Code)oe os CS. Wiemer and ©. them of the causes of PGS is necessary 26 jt often appears in otherwise healthy tortoises, Bucthermore, tortoises with signs of MBD like soft exrapace, lordsis, or extensive growth of the costal plates, relative wo the ental (lso named verebral plates often show n0 signs ‘of PGS. Although any explanation cannot be found concerning & possible influence of fenvionmental humidity in the scientific literature, private tortoise breeder experience suggests that PGS is rlated to the humidity under which the tortoises are kept (West, 1988). In view ofthe above, an experiment was designed to ascertain two possible causes of hhump formation in tortoises. This study was designed to esamine the effect of etary protein level and environmenisl humidity on the degree of pyramidal growth in the Satapaces, Information gained from the study should provide the bass fr advice to turtle Keepers in order to avon this problem in future Materials and methods Experimental animals Alrican spurred torwises (G. sleata) were selected for this experiment as PGS is common in this specie in captivity. [lathlings of African spurred tortoises also grow very fast. As the females ean produce cliches of more than 20 eggs, age and genetics can be controled in the hatchlings. Fifty hal-ibling tortoises were hatched athe Farm of a non-commercial ‘Austin breeder (parents: wo dames, one sire) ding late December to late January. They were selected from three clutches of two dames containing 21,29 and 27 haceilings. The hatchlings were placed into vansport hoxes immediately after hatching. Within 2 days of hatching they were transfered to teravia athe Institute of Notrtion atthe University of Veterinary Medicine, Vienna Experimental design and sample collection After exchading individals with deformations of the earspace or the horay plates at the breeding farm, the first $0 tortoises hatched were taken for the study group. At the University, the 50 tonoises were randomly distributed to five terraria and kept under identical conditions for 3 mantns except for dietary protein content and environmental humidity (Table 1). At the end of the feeding trial, the formed carapace humps were measured (Fig. 2) and blood samples were saken from the vena jugularis (to measure haematocrit, caleium and phosphorus levels. Weights were measured twice a month. Diet In each terrarium, water and sepia (squid-scelecon) calcium were supplied by three ceramic bowls (diameter 13 er) ad libitum, Every morning, the tortoises were fed a mixture of Table 1, Enviconmental humidity ud dietary protein concentestions for the five experimental groups of Grachelome deat Group A Group ® Group © Group D Group F. Environmental humidity 243-578 274-555 306-748 47.999 4599 | Feed Type! “Type? Type? Type? Typed Environmental humidiy: mean of eight weekly measured values af the maximum and minimum relsive Fumiciy in peecenage Material may be protected by copyright law (Title 17, U.S. Code) | Influence of Fig, 2. Measurement of tre Hid plates 2, 3 Table 2. Nutetional com) Dave Ens Crude prosin at rude fa 58 Sie tee Ba (mmg/l00 9) 240 Ca Pesto 44 dried pellets with defined au Germany) and endive sslad afterwards 2 1 until day 13 selectively, the sosked pelle ‘nerease feed aceepcance, Call (Table 2). This feeding progra secondary hyperparathyreidi technological reasons, values treatments, but this as nat balance differences by diffe. Weende analyses were used using an atomic ae Gezmany), and phosphorus ‘The teeraria consisted of gl 25 cm above the ground, Tw Front pane and at the eciling ‘each terrarium, the toztoisesin otherwise healthy tortoises ce, lordoss, or extensive growth Saal plates, often show eo signs acersing a possible ialueace of te tortoise breeder experience fh the tortoises are kepe (Weser, J ascertain two pomiblecaunes of o examine the effect of dietary ce of pyramidal growth in the Fide the bass for advice to turtle s experiment as PGS is common tortoises also grow very fast. As ye and genetics can be conctolled acthe farm of a non-commercial December to Tate January. They ag 21, 29 and 27 bacchlings. The ther hatching, Within 2 days of of Nutrition atthe University of collection apace or the horny plates at the for the study group. At the to five tervaria and kepe under rein content and envionmental e formed carapace humps w from the vena jugularis (0 Weights were measured twice & 0 were supplied by three ceramic tortoises were fed a mixture of srations for the five experimental np Group Group E | as 79.99 a5-08 ped Type2 Type fet of the maximum and minimum | Influence of bumiiy and det on pyramidal growth of carapace ° ‘ig, 2. Measurement ofthe H(ump) index. CICS: Central plates 1-5; H-index = (H-value (cone ples 2, 3) + ovale font’ plates 3, ]25 Fhvalue = a ‘Table 2 Nutritional composition of the dies fed (pereentage of dey matter, snalysed) “Type of feed To 7 | Days Days Days Days Days F108 106180115 samt 1S 16 150 | Grade protein. a 7 94 9 307 x07 Chace fe 58 52 “a “a Si St hide fibre be 1B ba 8 ha 19 CXlmyiong) = 22 23055571 2649) CaP rs an te ain dred pellets with defined nutrient comtents (prepared by ssniff Spevialdiaxen GmbH, Soest, Germany) and endive salad in the proportion of 1: 1 (pellets: endive) until day 105, afierwards 2: 1 until day 159 (all in feesh weight). To prevent che wutles from feeding selectively, the soaked pelles were mixed with chinly cut endive, Endive was used «0 ‘increase feed acceptance. Calcium and phosphorus were fed a 3 ratio of higher than 3:1 (Table 2). Ths feeding program was designed to prevent deficiencies and pathogeneses (eg secondaty hyperparathyroidism) which have been suggested as eauses of PGS, Due t0 technological ressons, values of ealcivm and the CaP ratio differed slightly beeween treatments, but this was not considered to be erucial, as the animals could theoretically balance differences by different sepia calcium consumption, ‘Weende analyses were used for the detection of crude nutrients. Caleium was analysed lusing an atomic absorption spectrophotomseter (3030 By Perkin Elmer, Uberlingen, Germany), and phosphorus was measured with 3 photometer. Environmental conditions Cages ‘The terraria consisted of glass (100 x 80x 80 em) with swo sliding panes at the front, 25 cm above the ground. ‘Two air slots for ventilation were located at the bottom of the front pane and at the ceiling pane. The substrate consisted of 4-5 em of bark humus. Tn exch terrarium, the tortoises could hie in caves made of bricks (30 x 65 x 10 em). Material may be protected by copyright law (Title 17, U.S. Code)7 CS. Wiemer and C. then Lighting Bach terrarium was illuminated by three different light sources. Under the top pane, a 150- W HQlamp (Teclumen, Castel Coffredo, Italy} was fixed, which supplied the animals with the necessary light invensity for activity. The coaaccted intermediate unit (containing the transformator) was located outside of the terraria, At the left side of the restarium ard about 25 em above the ground, x halogen hester lighted and heated che ares below. Also at this height, an ultvioles radiating fluorescent tube (UV-A and UV-B, Reptisun 5.23 Zoo Med Laboratories Inc, San Luis Obispo, CA, USA) was also provided at this height. The HQl-iamp was on 11h daily irom 8 4m to 7 pm. Heating light and UV-tube were activated for 3 h each morning (9-12 am) and 99 min each afternoon (3-4.50 pm). At the bottom of the central area of the terrarium 7609 lux, and under the halogen lamp, 14 000— 15 000 lux, were measured, respecsively. Haomidity control system Humidity was provided by plasic-bowls (40x40x15 em), filed with demineralized water with aromizers t0 produce fog. The bowls were located on top of the caves. Electric control units were used 10 regulace che atorizers, mintsining the level of relative humidity in the terraria. A connected sensor was locate inside the eaves. High humidity in terrarium D and E lead to humid soil The humidity values provided in Table | were measured directly tunder the top pane. Values measured inside the caves of the three humid terrariams were approximately 10-15% higher. Measurement of bump The wedge-shaped indentations between the second and he tied, and the third and fourth central plate of the earapace were measured. In this area, hump formation is most distines, ‘The Fi-value was defined asthe ratio of the depth (b) ofthe indentations at the growth-gap to the distance (a) berween two opposite edges of ateoks (Fig. 2). The mean of the two He values was considered to be the H{amp)-index to express the degree of hump formation of the tortoises. A. perfectly well-shaped carapace would have negative Hinde, due to negative depths (b) and consequently negative H-values Statsies To evaluate the differences berween ule groups with respect to H-indices, weight, Jhaematocrt, plasma esleium and plasma phosphorus appropriate non-paramecrcal tests of Kruskal-Wallis 1 ney Unest were used. The selecwed level of tests were conducted by using SPSS 9.9 (SPSS, Chicago, IL, USA). Results ‘The H-indices of gcoups B, C and D, raised under different levels of relative humidity but identical dietary protein concentration, were compared aad had formed distinetly different humps (Figs 3, 4-6; Table 3). Tortoises of group B (‘dry’) showed very prominent humps whereas the earapaces of tortoises of group E Chum!) were almost smooth, The humps of group C were between these two extremes. These three very different levels of hump development were significant (p< 0.001). Groups A and B both raised under dry conditions, but fed different levels of dietary protein, (B: 14.4/13.7 in % of dry matters. A: 19.4/19; Table 2) also produced different sized humps. Although the statistical significance probability of ercor (p = 9.06) of this difference was above the chosen level Material may be protected by copyright law (Title 17, U.S. Code) Infience of bam oe 26 Hinde 0. Fig, 3. This figure shows the dif: lkope under diferent levels of ela Aferent evs of dsxary pron (ged sre the median, Une 280h and 730% 9 outlier value. “" Valee it more { < 005), some influence of di ‘ised under humid conditions 30,7) showed no statistically pe first sight, the bigger African sp irmpression however elected on the humps were actully. meas ditfer among the groups but pl: higher than for groups B, C). 1 {group A). As ditary protein ct ‘weight of groups B, C, D (same ‘The combination of dry envieo induced by a nutritionally den tortoises ofthis study. Hemid ably. Lowering the level of dict development of PGS. Consider Ireratre regarding growth fi! they dilfer considerably from th reported his experience feo kes influence on hump formation. comiders some of the natural his have s highly variable ‘ood supp} ring a year and betwen yeasurces. Under the top pane, 4 150- xed, which supplied the animals ted intermediate unit (containing the left side of the terrarium and nd heated the atca below. Also at “A and UV-B, Reptisun 3.0; Zoo also provided at this height, The eating light and UV-tube were afternoon (3~4.30 pm). At the under the haloges lamp, 14 000 filed with demineralized water {r0p of the caves. Electric control level of relative humidity in che . High humidity in terarium D n Table 1 were measured dixectly the three humid terrariums were he chiral andl the third and fourth hun formation is most distinct w indentations atthe geovrtl gap Fig. 2). The mean of the two H- the degree of hump formation of lave 2 negative Flindex, due 10 respect 10 H-indices, weight, opriate non-paracnetrica tests of - used. The selecied level of ucted by using SPSS 9.0 (SPSS, at levels of relative humidity but < had formes distinctly diferente | showed very prominent huinps re almost smooth, The humps of c very different levels of hump and B both raised under dry L4/13,7 in % of dry matter vs. wimps. Although the statistical see was above the chosen level n Heindex & Group Fg 3. This igure shows he differences in hamp formation (Hindi) ofthe Sve groups of tries pe under dierent levee of rate humidity (group By Cand Dy Keunkl Wall tex) and fo {een levels of cary pron group A vB. group D ve; Mann-Whitney Utes Vals sbown bre the motion, the 25ch ana 75e porentes intergsare range) andthe highest and lowest non tier value "+" Value ie wove than LS fold and "value is more than feel! above te Inverqarle ange (p < 0.05), some influence of dietary protein is probable. Groups D and E, which were saised under humid conditions but fed different levels of dietary protein (D: 19.4/19 vs. E: 38.7) showed no statistically provable difference in hump size. It should be noted that at Ties sight, the bigger African spurred tortoises were believed to have bigger humps. That impression however refiected only the absolure hump size and was disproven later whee the humps were actually. measured. Regarding blood parameters, haematoerts did) not differ among the groups but plasma calcium levels did (Ca-level of group A, D, E was higher than for groups B, C). The P-levels were between 0.82 (group C) and 0.98 mmol/ {group A), As dietary protein comtent incressed body weight increased: however, the body “wcight of groups 8, C, D (same level of dietary protein) did nos di Discussion “The combination of dey environmental conditions and comparatively high growth rates induced by 2 nutritionally dense diet led to pysamidal groweh in the African spurred rontises of this study. Humid conditions suppressed the development of PGS consider ably. Lowering che level of dietary protein had a questionable suppressing effect on the development of PGS. Considering the published repors in the veterinary and herpetologic licentare regarding growth failure in chelonia, the results presented here are surprising a6 they differ considerably from the commonly propagated explanations. Only Weser (1988) reported his experience from keeping tortoises under humid condition and suspected sore influence on hump formation. However, the results presented are understandable if one considers some of the natural history of wild tortoises. Nearly all species of extant tortoises have & highly variable food supply both in quality and quantity of nutritional components during a year and between years. During humid seasons, tortoises are able to select from a Material may be protected by copyright law (Title 17, U.S. Code)n CS Wieser and C ben Influence of bai Table 3. Revues of the weight a Mean (N19) sD. & Mean (N19) 3 © Mean(N:10) | ‘sD. D ‘Mean (N : 10) sD. 5 ‘Mean (N : 10) sD. | ‘his abe shows the resus of the thre blood eases (haemasotiy e are not significant except the ue {grermediace prosein dst) va. ge Means of geoups not sharing’ variety of new groweh plants, cs rspi¢ growth, During very long 4 for tortoises are dry plants eis value. During these periods cori bur grow very little, Faster-grov above the sol, providing a higher tortoises live most of ther life hic fo on the ground surrounded by and habitats of torcoises in understandable that the abnorma level of protein may lead to heal very common both in privates prevalence of PGS in vorcoises regarding the pathological mee histological researeh, will it development of PGS and otker rickets or osteodystrophia bros could be that dehydration reduc ‘artiagineous tissue a the growie be caused by this low pressure becomes assfied and the collazes permanently. During each per higher humps, Bue this hypoches i intake or excretion of minerals the development of PGS. The supported or rejected such an ex TThe results presented indicat | cause of PGS, except that it 2) previously mentioned, faster gro Figs 4-6. Torsoises with thre levels of hump formation. Figores 4-6 show one vorteines from each of forming bigger humps. This group B (Gry environmeat), Ciaermediste omy) and wet envivonmest) tortoises produce bigger humps. level and degree of PGS, keeping be useful because of other prot from this study is tha, in order Material may be protected by copyright law (Title 17, U.S. Code) |Influence of buried diet on pyramidal growth of carapaces Table 3. Results of the weight gain ad blood values ofthe ive groups af seated tortoises Weidupng) Hke(%) Ca (mmolA) —_P (mmoi) a Mean (N : 10) We 192 336 295 sD 3 15 a4 B Mean (N : 10) i et as? ‘sD. A 08 os © Mean (N : 10) as ms ona ‘sD. 18 a7 D Mean (N : 10) Ea 165 O38 sD. 5 17 12 h Mean (N +19) ae 181 030 30. % 12 325 “This table shows he resus ofthe wh gain (ond weight minus tart weight of the tories) and thre Blood values (haematoer, caer snd phosphor sales). Direncesheeween the rope fe ot sipnieant cep the dora might gai of group A ow proscin dis) w= group D-DD {ieermediare protein die) vs. grnpf (igh protein ds) (Mann-Whitney Utes p< GOS) SESeas of groups a sharing aire ore ae signin diferent variety of new growth planss rchin autrtional contents corresponding to the period of rapid growth. Daring very long sesnns of dryaess and food scarcity, the main food source for tortoises are dry plants charactraed by « high fbre content and low overall nutritional value. During these periods rortois show litle ativity and probably osseous ccbnilding, but grow very litte. Faster-growig batchlings live under the cover of grass and directly above the soil, providing a higher lof environmental humidity. Finally, many species of torwoises live most of thei ite hidcescither in eaves (e.g. Teseuda harsfeldi, Gopheras sp.) for on the ground surrounded by high humidity even during arid conditions (for behaviour and habitats of corivises in nuns, see Ernst et al, 1994). Considering this, it is tunderstandable that che abnormal onbination of a dry environment and a persistent high level of protein may lead to health iarders, such as PGS. Such management conditions are very common both in private and insiutional situations, which prebably explains the high prevalence of PGS in tortoises is aprivity. Not until further information is available regarding the pathological meckaism of how low humidity leads ro PGS, such as histological research, will it be passible to evaluate any connection between the development of PGS and other dieses of the osseous tissue in turtles, such a MBD, rickets or osteodystropbia fibrosa. One possible explanation for the development of PGS could be thar dehydration reduces tie intracellular andl imercellular pressure on the soft cantlagineous tissue at the growing yp area A collapse of this tissue around the gap might be caused by this low pressure. Intoises are dehydrated for a longer period, the tissue becomes ossfied and the collapsed wleys’ between the central parts of the plates are Fixed permanently. During cach period oigowth the development of new valleys would create higher humps. Bus this hypothesis ads to be supported by further research. Differences in intake or excrctiog of minerals sha ealeium or phosphorus might also help co explain the development of PGS, The seats of blood assays, in the current stucy, neither supporced or rejected such an explsition “The results presented indicate thtthe level of dietary protein is probably not the main cause of PGS, except that it may predispose the tortoise to more rapid growth. AS previously mentioned, faster growig tortoises can provoke the erroneous impression of eG forming bigger humps. This mig explain the widespread belie thar faster growing al Dec eoumen ach tortoises produce bigger humps. Enz if there is a0 coanection between dietary protein level and degree of PCS, keeping theevel of dietary proven in tortoise diets low might sill seful because of other proteinsebted diseases, such as gour, The decisive conclusion from this study is tha, in order toveuce the incidence of PGS in growing tortoises, one Material may be protected by copyright law (Title 17, U.S. Code)7” CS Wiemer and © thew needs vo provide a sufficient level of envitonmental humigity. In particular, in onder vo keep the development af PGS ro a minimum, arcas with a relative humility of nearly 160% for hiding shovid be provided to the tortoises at all times. Acknowledgement We thank Prof Dr Howard 8. Hulan and Mag, Corel Gabler for their suppore and assistance. We slso. hive so. thank the World Tonle Association and the Avsirian Herpetologial Society (Gncereishisehe Geselashate ir Herpetelogie) for providing Snancil supporto ik ty References B1r0s, Ps Lawron, P. Cz Cooven J.C. 1997: Kompeninn der Repionbentien, Helen, ‘ingrnt, Theape. Silence, Haanover p ok aceasta, 0.200 Shrine de etalicen Pras Shines, Sige, p74 Fawr ©. Thy Lovion, Jy Dagnoon, R. Wy 1994 Twrler ofthe Unted Starr sod Canad SmishonanIntin Pres, Waringon Vann, p83. aon LF IDI: Boma ant Snel pes of Caoe Reptile Hasbanty. Kiger Pacing amp, Maar 8 Canucks Zan F995 Solem. In Gaon Ky ZAR Pe). Kroon der “eit, 3. Rule Selitche, mover, p. 73073 Hcmmen A.C, 198 Pct Encl f Reaping and Breding Tortie aed Hester “hanter, Caapace Dress, Loon, b.7 Jickson ©. Fs Coo fr 98: Notions den, In: CoOReR, J Es JnoMON E. F. (dh ‘Dace of be Repti Vol. 2 Academie Pes San Diego, pp. ease, W197 Die Landcirans Emap bls, Peg, Zach and Sas Meng Vers, ele pi Konten Gy 196: Krankhten de Ropilon wd Amphibion, Une, Setar 2.18 Meaaniun 5.1996: Vroinery Manegeme of ovis and Tar, Satkwel Sire, Oxford. ‘< Mao, D(A) 1986 Reptile Medicine and Sage W-B Saunders, Piladlphi, p39. Ons. Ja iran, W, 1972 De Landschderten Eurpas A Zensen, Witenberg Lathes. ‘8 Sanrarune, Ly 200: Shlnsenranbiite. bade Wering Rohmannsele, p36 Wausy | Gy WF Lendidhldeon, Oto, eget Zach ieTereim bale ‘Rehman, po. Wrsen Ry 198: Zur Focerilng be eho, Sara 10, 2-28, Wirctas y260 Die meitercane Lanse n der errcichen rai ns Wenn, (cd Nediemane Lanichderoen, Draco TorarisicThemenelt Ne 2 Jagan I Naor ond ‘Fee-Veran Minster py. 671 a Amtiore address C. §. Wessun and Coes, Instat for Emshrong, Ve Universe Wien, Veterinseplate 1, 1219 Vienna, Aust jrmediinisehe Material may be protected by copyright law (Title 17, U.S. Code)