New Species of the Genus Spintumix (Acari: Mesostigmata:Spinturnicidae) on Corynorhinus mexicanus (Chiroptera:

Vespertilionidae) in Central Mexico

JUAN B. MORALES-MALACARA AND RICARDO LOPEZ-W

Laboratorio de Acarologia, Departamento de Biologia, Facultad de Ciencias, Universidad Nacional Aut6noma de Mexico,Coyoacan 04510, Mexico, Distrito Federal

J. Med. Entomol. 35(4): 543-550 (1998)ABSTRACT The new species Spintumix traubi was found on the bat Corynorhinus mexicanus inTlaxcala, Mexico. The female, male, female deutonymph, male deutonymph, and protonymph aredescribed and illustrated.

KEY WORDS Spinturnicidae, Spintumix traubi, Corynorhinus mexicanus, bat ectoparasites

THE CENUS Spintumix comprises '"SO species world-wide (Uchikawa et al. 1994), and is mainly associatedwith vespertilionid bats. Only 7 species have beenrecorded from New World bats and only 4 from Mex-ico—S. americana (Banks, 1902); S. mexicana Rudnick,1960; S. orri Rudnick, 1960; and Spintumix sp. cited byMorales-Malacara and L6pez-W (1990), and Morales-Malacara (1996). The current description refers to theSpintumix sp. specimens mentioned in the 2 citedpapers. This new species is the 1st record of Spintumixon the American plecotine bat Corynorhinus mexica-nus Allen.

Materials and Methods

The Spintumix specimens were collected by R.L-W.or by R.L-W. and J.B.M-M. in Tlaxcala, Mexico (Mo-rales-Malacara and L6pez-W 1990) and originally allmites were deposited in the collection of J.B.M-M.Some of these mites, mounted in Hoyer's medium,were selected for the type series. Drawings were madewith a phase contrast Zeiss microscope with drawingtube. We have supplemented other information in thenew species description with morphological data oftaxonomic value according to Uchikawa et al. (1994),such as ornamentation of the dorsal, sternal, and ster-nogenital plates and the venter of the proximal legsegments and the topography of the proteronotal se-tae, among others. To use these characteristics, somespecimens were observed with phase contrast (PC) orNomarsky differential interference contrast (DIC)(Olympus Provis AX70 microscope). Other specimensthat had been preserved in 70% alcohol were preparedfor scanning electron microscopy (SEM) followingmethods given by Deunff (1982), and micrographs

1 liiiboratorio de Biologia y Ecologia de Mamfferos, Departamentodo Biologfa, Universidad Autonoma Metropolitana-Iztapalapa. Apar-tado Postal 55-535, Mexico 09340, Distrito Federal.

were made with a SEM microscope (JEOL JSM-5200).The nomenclature for idiosomal chaetotaxy is thatusedbyDomrow (1972) and Uchikawa (1990), and forsternal and gnathosomal chaetotaxy we follow Evans(1968). All measurements are in micrometers.

Following Domrow (1972) and Uchikawa et al.(1994), we have treated Spintumix as feminine ingender and species epithets accordingly.

Spintumix traubi Morales-Malacara & L6pez-W,new species(Figs. 1-18)

Female. Idiosoma broadly ovoid, longer than widewith a moderately enlarged opisthosoma. Dorsum(Fig. 1). Dorsal plate rhomboidal, longer than wide;anterior and posterior ends rounded; with 8 microsetalpairs and 5 pairs of pores, and with foveae large andnearly all divided into cells arranged in groups of1-3-2-4-2-2-1. Five pairs of long proteronotal setaelocated anterior and lateral to dorsal plate in groups of2-2-1; distance between pnl-pnl (19-27) < pnl-pn2(38-52), both anterior to dorsal plate. Three pairs ofhysteronotal setae: 1 pair of long poststigmal setae(pst) and 2 pairs of small caudal opisthosomal setae(cl-c2). Peritreme short, curving dorsad from stigmabetween coxae III and IV, continuing on dorsum overcoxa III, and coming back onto ventral surface to endbetween coxae II and III. Venter (Fig. 2). Tritosternumsmall, ribbonlike, wider than long. Sternal plate sub-pentagonal, slightly wider than long, with a regularpattern of scalelike ornamentation; sternal setae Stl-St3 small, on the border of plate; 2 pairs of pores (Figs.3,11). Genital area represented by a small transverseslit surrounded by radially striated cuticle, and with asmall semioval genital plate (Fig. 4); 1 pair of smallgenital setae posterior to genital area; 1 pair of metast-ernal setae laterally flanking genital area (Figs. 3 and4). 4 pairs of intercoxal IV setae; 2 pairs of medial

0022-2585/98/0543-0550302.00/0 © 1998 Entomological Society of America

544 JOURNAL OF MEDICAL ENTOMOLOGY Vol. 35, no. 4

Figs. 1 and 2. S. traubi n. sp., female. (1) Dorsum. (2) Venter.

opisthosomal setae, and 2 pairs of medium-size ven-trocaudal setae. Anal plate nearly round with 1 pair ofshort adanal setae, and a short postanal seta. Legs.Coxae I-IV robust; coxa I ornamented with slight (orimperceptible) striations; coxae II-IV distinctly orna-mented with striae; 1 curved stria intersected by 2slightly curved diagonal ridges particularly marked onposterodistal margin of coxa IV (Fig. 13); distal an-teroventral portion of femur I, genua I, and tibia I eachwith small shelllike striated plate; legs I greatly en-larged in width and more robust than other legs. Clawsof leg I enlaiged with small empodium. Setation of legsas in Table 1. Ventral leg setae small to medium sized,some serrated; except posterior seta of coxa II long,twice as long as anterior seta. Posterolateral setae ofgenu I, trochanter II, and femur II, and anterolateralsetae of femur IH-IV long, slender, and serrated. Pos-terolateral setae of trochanter I, femur I, genu II, andtibia I-II, and anterolateral setae of genu HI-IV, andtibia IH-IV small to medium sized and serrated. Dorsalleg setae short to long; all dorsal setae of genu I-IVlong, slender and serrated; both distal dorsal setae offemur I-IV long, slender, and serrated; femur I withproximal anterodorsal seta rather small, and proximalposterodorsal seta medium sized (Fig. 12); both prox-

imal dorsal setae of femur II small; femur IH-IV withproximal posterodorsal seta small. Seta md on tarsiII-IV medium in length and quite robust. Gnathosoma.The gnathosomal base is subrectangular in outline,with hypostomals 1 moderate in size, and hypostomals2-3 minute; gnathosomal setal pair moderate in sizebut larger than hypostomals 1; palpi slender with tro-chanter-tarsus chaetotaxy formula 1-5-6-9-9 + undi-vided apotele; palpal trochanter-femur with small andstout ventral setae with serrated tips (Fig. 5).

Measurements. HOLOTYPE 9 (followed in paren-theses by range and mean of 6 PARATYPE 9 9 [ex-ceptions noted]): Idiosoma length 1,000 (926-1,250,1,140), width at level of coxae H-IH 686 (686-876,793); dorsal plate length 578 (550-583,567), width 515(492-545, 514); sternal plate length 171 (155-171,166), width 211 (197-215,206); tritosternum length 18(16-29, 21), width 68 (57-76, 66). Setal lengths: pnl,98 (94-110,102 [n = 6]);pn2,96 (88-110,97 [n = 6]);pn3, 92 (80-96, 89); pn4,100 (95-110,102); pn5,103(97-103,99); pst, 104 (90-117,103); cl, 24 (21-34,27);c2, 27 (22-34, 26); Stl, 32 (24-32, 30 [n = 6]); St2, 27(25-29, 27 [n = 6]); St3, 24 (19-26, 23); genital setae29 (22-29, 25); ventrocaudal setae, internals 31 (28-37, 32) externals 33 (30-45, 37). Leg I, length (in-

July 1998 MORALES-MALACARA AND LOPEZ-W: S. traubi, NEW SPECIES 545

Figs. 3-5. S. traubi n. sp., female. (3) Sternal plate and genital area. (4) Genital plate. (5) Gnathosoma.

eluding coxa and pretarsus) 830 (704-870,819); femurI, width 196 (184-215,196).

Male. As in female except as noted. Smaller thanfemale. Idiosoma ovoid, longer than wide. Dorsum(Figs. 6 and 15). 2 pairs of long caudal opisthosomalsetae (cl-c2). Venter (Fig. 7). Tritosternum smallerthan in female, appearing as a small line, wider thanlong. Sternogenital plate flask-shaped, longer thanwide; anterior border essentially constituted by broadgenital opening; anterolateral borders of plate angular;lateral margins almost straight; posterior marginrounded; sternogenital setae Stl-St3 small on the bor-der of plate, with 2 pairs of pores, and with irregularpattern of glyph-shaped ornamentation (Figs. 8 and14). Metasternal and genital setae small, situated atalmost the same level posterior to sternogenital plate(Figs. 7 and 8). Intercoxa IV area with 6 pairs of shortand robust setae. Anal plate an inverted trapezoid withnearly rounded anterior angles, with terminal anusand 2 short adanal setae and 1 short dorso-caudalpostanal seta. Legs. Coxae I-IV robust, ornamentedwith more striae than in female; ventral surface offemora I-IV ornamented with striae; legs I lightlyrobust and longer than other legs, with normal clawsand similar to other legs. Gnathosoma. Spermadactylshort and curved (Fig. 16).

Measurements. ALLOTYPE 6 (followed in paren-theses by range and mean of 6 PARATYPE 86 [ex-

ceptions noted]): Idiosoma, length 721 (655-736,700), width 633 (546-634, 612 [n = 6]); dorsal platelength 574 (553-582, 567), width 478 (394-491, 464);Sternogenital plate, length 239 (230-270, 248), width194 (170-200, 188 [n = 5]); tritosternum length 9(3-9, 5), width 24 (23-42, 29). Setal lengths: pnl, 88(82-94, 89); pn2 80 (72-80, 76); pn3, 82 (69-84, 77);pn4, 84 (78-99, 88); pn5, 102 (85-102, 94); pst, 140(121-143,136); cl, 80 (74-85, 80); c2, 90 (81-90, 84);Stl, 34 (33-38,35 [n = 6]); St2,28 (20-33,26); St3,26(23-30, 25); genital setae 26 (23-29, 26 [n = 5]). LegI, length (including coxa and pretarsus) 866 (741-900,856); femur I, width 150 (142-144, 150).

Female Deutonymph. As in adult female except asnoted. Smaller than female. Dorsum. Dorsal plate withcelled foveae as in adults, but few discernible. Opis-thosomal setae (cl-c2) as in male. Peritreme longerthan in adults, beginning with the stigma near to ven-tral portion of idiosoma between coxae III and IV,continuing dorsad at level of anterior portion of coxaIV and turned forward encompassing coxa III andending on ventral surface between coxae II and III.Venter (Fig. 17). Sternal plate flask-shaped with an-terior border narrow and truncated, longer than wide,with sternal setae Stl-St3 small on border of plate, and2 pairs of pores; sternal plate ornamentation as in male(Figs. 9 and 17). Genital and metasternal setae small,both situated almost at same level, posterior to sternal

Table 1.

11IIIIV

Leg eliaetotaxy of femnle, male,

Coxa

2221

Trochanter

0-0/4-1(5)1-0/3-1(5)1-1/3-0(5)1-1/3-0(5)

and deiitonyniphs of S.

Femur

2-4/3-2(11)1-4/3-1(9)1-3/2-0(6)1-3/2-0(6)

traubi

Genu

1-4/3-1(9)1-4/2-1(8)1-4/2-1(8)1-4/2-1(8)

Tibia

1-4/3-1(9)1-3/2-1(7)1-3/2-0(6)1-3/2-0(6)

Tarsus

5-18/5-5(33)3-5/5-3(16)3-5/5-3(16)3-5/5-3(16)

546 JOURNAL OF MEDICAL ENTOMOLOGY Vol. 35, no. 4

8r i\

Figs. 6-8. S. traubi n. sp., male. (6) Dorsum. (7) Venter. (8) Sternogenital plate.

plate. Intercoxa IV area with 6 pairs of small setae, and2 pairs of long ventrocaudal setae (Fig. 9). Anal platenearly square-rounded, with terminal anus and 2 shortadanal setae and 1 short dorso-caudal postanal seta.Legs. Coxae I-IV robusts, ornamented with striae as inmale; 1 curved stria with 2 diagonal ridges particularlymarked on posterodistal margin of coxa IV as in adults(Fig. 17); ventral surface of femora I—II ornamented asin male; legs I ligthly robust and longer than other legs,with normal claws and similar to other legs. Setationof legs as in adults (Table 1); except posterodistal setaof coxa II slightly longer (never twice) than antero-distal seta.

Measurements. PARATYPE DN9 (followed in pa-rentheses by range and mean of 6 PARATYPE DNN 9[exceptions noted]): Idiosoma, length 719 (683-770,723), width 634 (634-674,656); dorsal plate length 580(574-630, 598), width 550 (550-595, 569); sternalplate, length 220 (214-223, 218), width 197 (173-197,180); tritosternum length 21 (19-26, 23), width 43

(43-60,51). Setal lengths: pnl, 117 (117-125,121 [n =6]); pn2, 102 (94-113, 106); pn3, 94 (94-113, 104);pn4,102 (102-114,109); pn5,110 (106-123,116); pst,152 (143-154,149); cl, 102 (102-114,107 [n = 6]); c2,127 (117-129,123 [n = 6]); Stl, 27 (25-31,27); St2,23(21-24, 23); St3, 23 (19-23, 21); ventrocaudal setae,internals 74 (73-80, 75 [n = 6]) externals 76 (65-76,70 [n = 6]). Leg I, length (including coxa and pre-tarsus) 857 (731-857, 795 [n = 4]); femur I, width 170(170-211, 192).

Male Deutonymph. As in female deutonymph ex-cept as noted. Smaller than female deutonymph. Ven-ter (Fig. 10). Intercoxa IV area with 6 pairs of smallsetae and without ventrocaudal setae.

Measurements. PARATYPE DNo* (followed in pa-rentheses by range and mean of 5 PARATYPE DNNd[exceptions noted]): Idiosoma, length 657 (633-686,660), width 584 (584-637,602); dorsal plate length 555(543-559, 552), width 486 (480-506, 491); sternalplate, length 200 (200-222, 209), width 159 (159-182,

July 1998 MORALES-MALACARA AND L 6 P E Z - W : S. traubi, NEW SPECIES 547

Figs. 9-10. S. traubi n. sp.; sternal plate, intercoxa IV area, coxae IV, anal plate and ventro-caudal margin. (9) Femaledeutonymph. (10) Male deutonymph.

175); tritosternum length 22 (19-23, 21), width 51(51-61, 57). Setal lengths: pnl, 106 (102-116, 106);pn2, 91 (87-109, 96); pn3, 86 (86-98, 96); pn4, 98(94-108,100);pn5,101 (98-117,104 [n = 5]);pst, 140(137-150,143); cl, 92 (77-106, 91); c2,107 (102-119,109); Stl, 29 (26-29, 28); St2, 23 (20-24, 22); St3, 22(21-23, 22). Leg I, length (including coxa and pretar-sus) 797 (797-838, 821 [n = 3]); femur I, width 167(167-188,175).

Protonymph. As in deutonymphs except as noted.Smaller than female deutonymph. Dorsum. Dorsalplate rhomboidal, longer than wide, with 8 microsetalpairs and 3 pairs of pores, and with 10 or 11 smallcentral foveae indistinctly. Peritreme as in adults. Ven-ter (Fig. 18). Sternal plate flask-shaped with anteriorborder narrow, and truncated but broader thandeutonymphs; sternal plate ornamentations with pat-tern of irregular broad lines, curves, points, and someglyphs (as precursors of glyph-shaped ornamentations

in subsequent stage) (Fig. 18). Intercoxa IV area with4 pairs of small setae, lacking ventrocaudal setae as inmale deutonymph. Anal plate nearly squared-rounded, with 2 short adanal setae, and 1 short dorso-caudal postanal seta. Legs. Coxae I-IV robust, orna-mented with striae; curved stria and diagonal ridges onposterodistal margin of coxa IV not discernible, onlyobserved undulated lines (Fig. 18); ventral surface offemora I-IV ornamented with striae as in male andmale deutonymph. Setation of legs as in Table 2; pos-terolateral setae of legs I—II, anterolateral setae of legsII-IV, dorsal setae of femur I-IV, and dorsal setae ofgenu I-IV similar in proportions to adults anddeutonymphs, but quite small; except posterodistaland anterodistal setae of coxa II similar in size. Gna-thosoma. Palpi trochanter-tarsus chaetotaxy formula1-4-5-8-9 + undivided apotele.

Measurements. PARATYPE PN (followed in paren-theses by range and mean of 6 PARATYPE PNN [ex-

JOURNAL OF MEDICAL ENTOMOLOGY Vol. 35, no. 4

Figs. 11-18. S. fraufot n. sp. (11) Female sternal plate (SEM). (12) Female femur I dorsal view (SEM). (13) Female coxaIV (PC). (14) Male sternogenital plate (SEM). (15) Male dorsum (DCI). (16) Male spermadactyl (PC). (17) Femaledeutonymph venter (PC). (18) Protonymph venter (PC).

July 1998 MORALES-MALACARA AND LOPEZ-W: S. traubi, NEW SPECIES 549

Table 2.

IIIIIIV

Leg rliaelotaxy of protonympli of S. traubi

Coxa

2221

Trochanter

0-0/3-1(4)1-0/2-1(4)1-1/2-0(4)1-1/2-0(4)

Femur

2-4/2-2(10)1-4/2-1(8)1-3/1-0(5)1-3/0-0(4)

Genu

1-4/2-1(8)1-4/1-1(7)1-4/1-1(7)1-4/2-0(7)

Tibia

1-4/2-1(8)1-3/2-1(7)1-3/2-0(6)1-3/2-0(6)

Tarsus

5-15/6-3(27)3-5/4-3(15)3-5/4-3(15)3-5/4-3(15)

ceptions noted]): Idiosoma, length 660 (643-699,668), width 598 (555-617,587); dorsal plate length 558(521-558, 539), width 486 (463-486, 476); sternalplate, length 202 (187-210,197), width 161 (155-184,166); tritosternum length 20 (16-24, 21), width 53(51-55, 53). Setal lengths: pnl, 116 (106-127, 116);pn2,115 (106-115,109); pn3,110 (96-110,105); pn4,114 (106-120, 115); pn5, 99 (98-108, 102); pst, 130(118-130,124); cl, 78 (73-94,83); c2,94 (90-104,96);Stl, 29 (26-29, 28); St2, 25 (21-25,23); St3,24 (22-24,23 [n = 5]). Leg I, length (including coxa and pre-tarsus) 736 (690-807, 759); femur I, width 157 (153-184, 159).

Type Series. HOLOTYPE 9, 4 PARATYPE $ 9 , 3PARATYPE 66,5PARATYPEDNN9,6PARATYPEDNNd, and 1 PARATYPE PN, ex Corynorhinw mexi-canw 9, El Tunel 10 km E, Tlaxco, Tlaxcala, Mexico,5-IH-83, R. L6pez-W. 1 PARATYPE 6, ex C. mexica-nus, same data, 13-VIH-82, R. L6pez-W (UAMI-4777).2 PARATYPE PNN, ex C. mexicanw 6, same data,20-XI-82, R. L6pez-W. 1 PARATYPE 9,2 PARATYPE66, and 1 PARATYPE PN, ex C. mexicanw 6, samedata, 20-1-83, R. L6pez-W (UAMI-4805). 1 PARA-TYPE 9, ex C. mexicanw, same data, 27-XI-83, R.Lopez-W & J.B. Morales-Malacara. 1 PARATYPEDN9, ex C. mexicamis, same data, 17-HI-84, R.L6pez-W & J.B. Morales-Malacara. ALLOTYPE 6, 1PARATYPE DN9, 3 PARATYPE PNN, ex C. mexica-nw, same data, 27-1-85, R. L6pez-W & J. B. Morales-Malacara.

The holotype, allotype, 1 paratype 9,1 paratype 6,2 paratype DNN 9,1 paratype DN6, and 2 paratypePNN are deposited in the Colecci6n Nacional de Ac-aros (CNAC) at the Instituto de Biologfa, UniversidadNacional Autonoma de Mexico. The remaining para-types are in the Morales-Malacara Collection ( 1 9 , 16,1DN 9,1DN6",1PN); National Museum of NaturalHistory, Smithsonian Institution, Washington, D.C. (19 ,1 6,1 D N 9 , 1 DNo*, 1 PN); B.P. Bishop Museum,Honolulu, HI (1 9,16,1 DN9, 1 DN6, 1 PN);American Museum of Natural History, New York ( 1 9 ,1 6,1 DN 9,1 DN6,1 PN); Natural History Museum,London (1 9, 1 6, 1 DN9, 1 DNo*, 1 PN).

Etymology. The species name is dedicated to thememory of Dr. Robert Traub for his outstanding workon parasitology.

Discussion

Some features of S. traubi are similar to those of S.americana (Banks, 1902), including few hysteronotalsetae and tiny dorsal setae on femora I-IV (Rudnick

1960). A preliminary analysis of S. americana fromUruguay (ex Myotis) and Mexico (ex Myotis velifer),also indicates similar topography of proteronotal se-tae, and ornamentation of proximal leg segments, par-ticularly on coxa IV (compare, Fig. 13). These mor-phological similitudes confirm the opinion ofUchikawa et al. (1994) that S. americana does notbelong to the S. myoti species group. Further taxo-nomic evaluation of all Spinturnix species from theAmericas is needed to define species groups for theNew World spinturnicid fauna.

However, S. traubi is separated from S. americanaand from other known species in its genus by havingthe largest leg I with large claws and small empodium;the presence of a distal anteroventral striated shell-like plate on femur I, genua I, and tibia I in the female;and glyph-shaped ornamentation on the sternogenitalplate of male and the sternal plate of thedeutonymphs.

Spinturnix traubi is a Neartic species, as is its hostCorynorhinus mexicanw (Morales-Malacara andL6pez-W 1990), and represents the 4th species in thegenus recorded from Mexico and the 7th from theNew World. As we have noted (Morales-Malacara andL6pez-W 1990), this is the 1st record of a Spinturnixspecies on an American bat in the tribe Plecotini. It isnoteworthy that the Old World plecotine bats have awell-defined group of Spinturnix parasites, the S. ple-cotina group (Uchikawa et al. 1994). The morphologyof S. traubi indicates that this species is not related tothe S. plecotina group and that it is closely related toS. americana. On the other hand, no Plecotini batsfrom the New World have yet been shown to havespinturnicids, with the exception of C. mexicanw (Mo-rales-Malacara and L6pez-W 1990). This fact may sup-port the most recent phylogenetic analyses, proposedby Tumlison and Douglas (1992), and Bogdanowiczand Owen (1996), in which the Plecotini are separatedinto 2 groups—the Old World species in the genusPlecotus Geoffroy St.-Hilaire, and the New World spe-cies of the Nearctic Region in the genus CorynorhinusAllen. Apparently there is a close relationship of thehosts of S. traubi and S. americana as shown by cyto-genetic studies of vespertilionid bats of the tribes Ple-cotini and Myotini (Bickham 1979, Fedyk and Rupre-cht 1983, Stock 1983, Qumsiyeh and Bickham 1993).Also, it is well known that C. mexicanw and Myotisvelifer frequently share the same caves or refuges(Morales-Malacara and L6pez-W 1990). Therefore S.traubi and S. americana may have evolved from thesame lineage.

550 JOURNAL OF MEDICAL ENTOMOLOGY Vol. 35, no. 4

Acknowledgments

The senior author expresses his appreciation to VictorCastaiio (Intituto de Ffsica, Universidad Nacional Aut6nomade M6xico) and Martha Corona (Laboratorio de Acarologfa,Facultad de Ciencias, Universidad Nacional Aut6noma deM6xico) for providing facilities for scanning electron micro-scope micrographs. We also thank Jos6 Antonio Hernandezand Homer A. Gamboa (Laboratorio de Microcine, Facultadde Ciencias, Universidad Nacional Aut6noma de M6xico) forproviding phase contrast and differential interference con-trast microscope micrographs. For review of the manuscript,we express our appreciation to Anita Hoffmann (Laboratoriode Acarologfa, Facultad de Ciencias, Universidad NacionalAutdnoma de M6xico), Tila M. P6rez (Instituto de Biologfa,Universidad Nacional Aut6noma de Mexico), GerardoPeYez-Ponce de Le6n (Instituto de Biologfa, UniversidadNacional Aut6noma de M6xico), and Oscar Flores (Museo deZoologfa, Facultad de Ciencias, Universidad Nacional Au-t6noma de Mexico). Financial assistance was provided byDirecci6n General de Asuntos del Personal Acad6mico, Uni-versidad Nacional Autdnoma de Mexico, Grant No.IN215796.

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Received for publication 31 July 1997; accepted 18 De-cember 1997.