Cyrtodactylus panitvongi, Pauwels & Chotjuckdikul & Donbundit & Sumontha & Meesook, 2024
publication ID |
https://doi.org/ 10.11646/zootaxa.5512.3.3 |
publication LSID |
lsid:zoobank.org:pub:337DCC9D-2FC7-4006-B1E9-B2AF308475CD |
DOI |
https://doi.org/10.5281/zenodo.13861994 |
persistent identifier |
https://treatment.plazi.org/id/03998786-FFE2-FB2C-45C7-71D66AA60414 |
treatment provided by |
Plazi |
scientific name |
Cyrtodactylus panitvongi |
status |
sp. nov. |
Cyrtodactylus panitvongi sp. nov.
( Figures 1–6 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE 5 View FIGURE 6 )
Holotype. CUMZ-R-2596 (field no. MS 757 ), adult male caught on 27 December 2022 in Tham (= Cave) Khao Chan , Wat (= Temple) Tham Khao Chan (14°58’42.2”N, 101°18’40.6”E), Tha Luang District , Lopburi Province, central Thailand, by Nonn Panitvong , Natthaphat Chotjuckdikul , Saranon Charoensuk, Natthasuda Donbundit and Montri Sumontha. GoogleMaps
Paratypes (3). CUMZ-R-2597 (field no. MS 758 ), adult male, and CUMZ-R-2594–2595 (field nos. MS 759 and MS 760 , respectively), adult females; same locality, date and collectors as for the holotype .
Diagnosis. Cyrtodactylus panitvongi sp. nov. can be distinguished from all other congeneric species by the combination of its maximal known SVL of 85 mm, 15–18 interorbital scales, 35–37 interciliary scales, 10–12 supralabials, 9–11 infralabials, 29–33 paravertebral tubercles of which 22–25 between limb insertions, 18 longitudinal rows of dorsal tubercles at midbody, 40–42 ventral scale rows at midbody, poorly defined, non-denticulate ventrolateral folds, a series of 13 or 14 enlarged femoroprecloacal scales including 2 pores on the proximalmost scales on each side separated by a single smaller medial scale in males, or six precloacal pits on a continuous series of 32–34 enlarged femoroprecloacal scales in females, no precloacal groove or depression, two or three postcloacal spurs on each side of the base of the tail in males and females, no interdigital webbing, 16–19 total subdigital lamellae on 4th toe, a row of widened median subcaudal plates, a banded dorsal pattern including an uninterrupted nuchal collar and three dark bands on dorsum, 10 or 11 dark-colored rings on original tail, and a golden iris.
Description of the holotype. Adult male ( Figures 1 View FIGURE 1 , 2 View FIGURE 2 & 5 View FIGURE 5 ). SVL 74.0 mm. TailL 93.3 mm, tail original and complete. Head relatively long (HeadL/SVL ratio 0.30), wide (HeadW/HeadL ratio 0.66), not markedly depressed (HeadD/HeadL ratio 0.43), distinct from slender neck. Loreal region inflated, canthus rostralis not prominent. Snout elongate (SnOrb/HeadL ratio 0.41), rounded, longer than orbit diameter (OrbD/SnOrb ratio 0.85); scales on snout small, rounded to oval, granular, mostly homogeneous, larger than those on crown, interorbital and occipital regions. Eye large (OrbD/HeadL 0.35); pupil vertical with crenelated margins; supraciliaries short, bearing small conical spines. Ear opening rounded, small (EarL/HeadL 0.04); orbit to ear distance smaller than orbit diameter (OrbEar/OrbD 0.81). Rostral much wider (3.8 mm) than deep (2.0 mm), rostral crease about half of rostral height. Two enlarged supranasals separated from one another by two small superposed scales. Rostral in contact with first supralabials, nostrils, supranasals and internasal. Nostrils oval, more or less laterally directed, each surrounded by supranasal, rostral, first supralabial and three postnasals. Three or four rows of small scales separate orbit from supralabials. Mental triangular, distinctly wider (2.9 mm) than deep (1.8 mm). A single pair of greatly enlarged postmentals in broad contact behind mental, each bordered anteromedially by mental, anterolaterally by first infralabial, posterolaterally by an enlarged lateral chinshield; both postmentals collectively bordered posteriorly by three gular scales. Supralabials to mid-orbital position 8/7, enlarged supralabials to angle of jaws 10/11. Infralabials 11/10. Interorbital scale rows across narrowest point of frontal bone 18.
Body slender, moderately short (AG/SVL 0.39; TrunkL 43.5 mm) with poorly defined, non-denticulate ventrolateral folds. Dorsal scales relatively homogeneous, domed; dorsal tubercles about five times size of adjacent dorsal scales, extending from nape onto tail, smaller tubercles on postocular region, crown, occiput and nape; most tubercles bearing a strong keel, less marked on lower flank tubercles; tubercles on posterior trunk and sacral region most prominent; tubercles in 18 rows at midbody, separated from one another by two to four, but generally three, dorsal granules. Ten or 11 granular dorsal scales surround each tubercle. Ventral scales larger than dorsals, smooth, oval and subimbricate, largest on posterior abdomen and in precloacal region. Midbody scale rows across belly between ventrolateral folds 42. Gular region with homogeneous, smooth, juxtaposed granular scales. No precloacal groove or depression. On each side a continuous series of 13 enlarged femoroprecloacal scales, the two proximalmost bearing each a pore; the two series separated by a diastema of one poreless medial scale. Postcloacal spurs each bearing two enlarged, conical scales.
Scales on palm and sole smooth, rounded to oval or hexagonal, slightly domed. Scalation on dorsal surface of hind and forelimbs similar to body dorsum with enlarged tubercles interspersed among smaller scales. Fore- and hind limbs relatively long, slender (FaL/SVL 0.15, TibL/SVL 0.21). Digits long, slender, inflected at interphalangeal joints, all bearing robust, slightly recurved claws. Basal subdigital lamellae broad, oval to rectangular, without scansorial surfaces (5-4-4-5-6 right manus, 3-4-5-6-6 right pes); narrow lamellae distal to digital inflection and not including ventral claw sheath: 10-12-12-10-11 (right manus), 10-11-13-12-15 (right pes); no interdigital webbing. Relative lengths of digits: IV>III>V>II>I (manus), IV>V>III>II>I (pes). Tail original and complete, gently tapering to pointed tip, distinctly longer than SVL (TailL/SVL ratio 1.26). A single median row of enlarged subcaudal plates.
In life, dorsal ground color of head, dorsum and limbs light olive brown. Poorly marked reticulations on head. Area above orbits bluish. A brown continuous nuchal loop, pursuing a brown preorbital stripe, connects orbits. Three brown bands on dorsum, longer than their light interspace, the anterior one beginning above the shoulder; the anterior and posterior borders of the collar and these brown bands black; numerous black irregular marks on the light interspaces ( Figure 1 View FIGURE 1 ). Irregular thin, black bars on upper surface of limbs, hands and feet. Background color of anterior part of tail olive brown as dorsum, turning to white posteriorly, with ten dark rings (dark brown anteriorly, turning to black posteriorly), encircling the tail; irregular black marks on the light interspaces between the dark rings. Iris golden. Supralabials, infralabials, throat, venter and undersides of fore- and hind limbs uniformly whitish.
Variation. Main morphometric and meristic characters of the type series are provided in Table 1 View TABLE 1 . Similarly to the holotype, ten or 11 granular dorsal scales surround each tubercle in the paratypes. While both males show four precloacal pores separated by a diastema of one scale, both females have a continuous series of six pitted precloacal scales, and more enlarged femoroprecloacal scales (32–34 versus 26–28). Besides the holotype, only one other type, the female CUMZ-R-2595 , has a complete, original tail, with a TailL/SVL ratio of 1.08, thus proportionally a bit shorter, and ten dark rings ( Figure 5 View FIGURE 5 ). The reticulate pattern on the head is more marked on two paratypes ( CUMZ-R-2594 and CUMZ-R-2597 ; Figures 3 View FIGURE 3 and 4 View FIGURE 4 ) than in the holotype. The only known juvenile shows a pattern similar to that of the adults, but more contrasted, with 11 dark rings on tail ( Figure 6 View FIGURE 6 ). The regenerated parts of tail show no more rings, but a light to dark brown background color mottled with black, above and below, and no more widened subcaudals.
Distribution and natural history. All adult individuals observed, including the type-series, were found at night within the twilight zone and the entrance zone of Khao Chan Cave or just outside its entrance. Several juveniles were observed, all just outside the cave. Cyrtodactylus panitvongi sp. nov. is a trogloxene species (sensu Jantarit & Ellis 2023) with a karst ecotype (sensu Grismer et al. 2020b, 2021c). It is so far known only from its type-locality ( Figures 7 View FIGURE 7 and 8 View FIGURE 8 ). Within the cave, near its entrance, it was observed with the reptiles Calotes versicolor (Daudin) ( Agamidae ), Dixonius siamensis (Boulenger) , Gehyra wongchan Pauwels, Meesook, Kunya, Donbundit & Sumontha , Gekko gecko (Linnaeus) , G. pradapdao Meesook, Sumontha, Donbundit & Pauwels , Hemidactylus frenatus Duméril & Bibron , H. platyurus (Schneider) ( Gekkonidae ), Lycodon capucinus Boie , L. davisonii (Blanford) ( Colubridae ) and Trimeresurus macrops Kramer ( Viperidae ), and the amphibians Duttaphrynus melanostictus (Schneider) ( Bufonidae ) and Polypedates megacephalus Hallowell ( Rhacophoridae ). Just outside the cave entrance, it was found with the amphibians Microhyla heymonsi Vogt ( Microhylidae ) and Fejervarya limnocharis (Gravenhorst) ( Ranidae ).
Etymology. The specific epithet honors our friend, the Thai zoologist, conservationist and photographer Nonn Panitvong, for his contributions to the knowledge of the fauna of the Kingdom of Thailand. We suggest the following common names: ตุ๊กกายถ้ำลพบุรี (Took-kai-tham-Lopburi; Thai), Lopburi Bent-toed Gecko (English) , and Cyrtodactyle de Lopburi (French).
Comparison to other species. Cyrtodactylus panitvongi sp. nov. is readily distinguishable from C. cardamomensis Murdoch, Grismer, Wood, Neang, Poyarkov, Ngo, Nazarov, Aowphol, Pauwels, Nguyen & Grismer , C. intermedius Smith , C. kulenensis Grismer, Geissler, Neang, Hartmann, Wagner & Poyarkov , C. thylacodactylus Murdoch, Grismer, Wood, Neang, Poyarkov, Ngo, Nazarov, Aowphol, Pauwels, Nguyen & Grismer and other members of the C. intermedius complex by the presence of numerous black marks on the light interspaces on dorsum and tail (versus absence of black marks), and by the absence (versus presence) of a dotted line of white tubercles outlining the dark dorsal bands (see Murdoch et al. 2019 and Grismer et al. 2020a, 2021a, 2023a). By its discontinuous series of 4 PrePo in males, it is moreover distinguished from Cyrtodactylus cardamomensis (9 or 10 PrePo in a continuous series in males), C. intermedius (8–10 PrePo in a continuous series in males), C. kulenensis (9 or 10 PrePo in a continuous series in males) and C. thylacodactylus (7–9 PrePo in a continuous series in males).
By its regular banded dorsal pattern and its four precloacal pores in an interrupted series in males, Cyrtodactylus panitvongi sp. nov. is easily recognizable from C. amphipetraeus Chomdej, Suwannapoom, Pawangkhanant, Pradit, Nazarov, Grismer & Poyarkov (very irregular bands or blotches; 10–12 FemPo and seven to nine PrePo in males; see Chomdej et al. 2020), C. angularis (Smith) (blotched; five or six PrePo pores in males; see Smith 1921 and Ulber & Grossmann 1991), C. jarujini (blotched; 52–54 FemPrePo in males; see Ulber 1993; Chan-ard 2007; Sumontha et al. 2008 and Thongnetr et al. 2021), C. monilatus Yodthong, Rujirawan, Stuart, Grismer, Aksornneam, Termprayoon, Ampai & Aowphol (blotched; absence of pores in both sexes; see Yodthong et al. 2022), C. oldhami (Theobald) (striped/dotted) and C. phetchaburiensis Pauwels, Sumontha & Bauer (striped/blotched; five PrePo in males; see Theobald 1876 and Pauwels et al. 2016a).
It differs from Cyrtodactylus lenya Mulcahy, Thura & Zug by its possession of three (versus five) dark bands on dorsum and of precloacal pores (versus absence of pores) (see Connette et al. 2017). Its nuchal collar and dorsal bands are much longer than those of Cyrtodactylus saiyok Panitvong, Sumontha, Tunprasert & Pauwels , whose males show five PrePo in a continuous series (see Panitvong et al. 2014 and Aksornneam et al. 2023). It differs from Cyrtodactylus sumonthai Bauer, Pauwels & Chanhome by its numerous black marks on the light interspaces on dorsum (versus a few dark marks), its three dark bands on dorsum (versus 4). Cyrtodactylus sumonthai has one or two PrePo and no enlarged femoral scales (see Bauer et al. 2002 and Panitvong et al. 2012).
Cyrtodactylus panitvongi sp. nov. does not have a prehensile tail, contrary to C. denticulatus Chomdej, Suwannapoom, Pradit, Phupanbai & Grismer View in CoL (13 PrePo in males), C. fluvicavus Grismer, Aowphol, Yodthong, Ampai, Termprayoon, Aksornneam & Rujirawan View in CoL (14 or 15 PrePo and 8–11 FemPo in males), C. interdigitalis Ulber View in CoL (14 PrePo and 17 FemPo in males), C. rivularis Grismer, Aowphol, Yodthong, Ampai, Termprayoon, Aksornneam & Rujirawan View in CoL (series of enlarged precloacal and femoral scales disjunct), C. rukhadeva Grismer, Suwannapoom, Pawangkhanant, Nazarov, Yushchenko, Naiduangchan, Le, Luu & Poyarkov View in CoL (17 PrePo and 20 FemPo in males), C. thongphaphumensis Grismer, Rujirawan, Chomdej, Suwannapoom, Yodthong, Aksornneam & Aowphol View in CoL (15 PrePo and 12–16 FemPo in males) and C. uthaiensis Grismer, Aowphol, Yodthong, Ampai, Termprayoon, Aksornneam & Rujirawan View in CoL (14 PrePo and 12 FemPo in males) (see Grismer et al. 2021b, 2022a, 2023b and Chomdej et al. 2023). Cyrtodactylus panitvongi sp. nov. lacks the webbing and the strongly developed ventrolateral folds of C. ngati Le, Sitthivong, Tran, Grismer, Nguyen, Le, Ziegler & Luu View in CoL (13 PrePo and 14 FemPo in males), and has a much more contrasted dorsal pattern (see Le et al. 2021 and Grismer et al. 2022b).
Cyrtodactylus panitvongi sp. nov. has a longer nuchal collar than C. auribalteatus Sumontha, Panitvong & Deein View in CoL (six PrePo and nine or ten FemPo), and the adults lack the round spots within the light dorsal bands of the latter (see Sumontha et al. 2010). Contrary to Cyrtodactylus panitvongi sp. nov., the light dorsal bands of C. samroiyot Pauwels & Sumontha View in CoL (seven PrePo in a continuous series in males) and C. tigroides Bauer, Sumontha & Pauwels View in CoL are uniform or with a few round spots (see Bauer et al. 2003 and Pauwels & Sumontha 2014). While the light bands of Cyrtodactylus panitvongi sp. nov. bear irregular black marks, those of C. chanhomeae Bauer, Sumontha & Pauwels View in CoL show a straight band (32–34 FemPrePo in males; see Bauer et al. 2003).
Within a 400 km radius, the only species showing patterns resembling that of Cyrtodactylus panitvongi sp. nov. are C. kunyai Pauwels, Sumontha, Keeratikiat & Phanamphon View in CoL , C. muangfuangensis Sitthivong, Luu, Ha, Nguyen, Le & Ziegler View in CoL and C. papilionoides Ulber & Grossmann. View in CoL Cyrtodactylus panitvongi sp. nov. is distinguished from C. kunyai View in CoL by its uninterrupted (versus interrupted) nuchal collar, a golden (versus dark orangeish) iris, distinctly higher VentR (40–42 versus 34), and absence (versus presence) of FemPo in males (see Pauwels et al. 2014). It differs from C. muangfuangensis View in CoL by its three (versus four) dark bands on dorsum, its higher DTR (18 versus 15 or 16), higher VentR (40–42 versus 31–37), lower number of PrePo (four versus six), absence of FemPo (versus presence), and absence of pores in females (versus presence) (see Sitthivong et al. 2019). It can be separated from C. papilionoides View in CoL based on its uninterrupted (versus partly or totally interrupted) nuchal collar, its higher DTR (18 versus 12–14), distinctly higher VentR (40–42 versus 30–34) and interrupted (versus uninterrupted) series of PrePo.
Holotype, CUMZ-R-2596 | Paratype, CUMZ-R-2597 | Paratype, CUMZ-R-2594 | Paratype, CUMZ-R-2595 | |
---|---|---|---|---|
Sex | Male | Male | Female | Female |
SVL | 74.0 | 77.0 | 85.0 | 84.4 |
TrunkL | 43.5 | 45.7 | 52.6 | 49.0 |
AG | 29.0 | 35.4 | 38.4 | 35.3 |
TailL | 93.3 (original) | 76.0 (68.3 regenerated) | 88.5 (63.6 regenerated) | 91.4 (original) |
TailW | 7.3 | 7.2 | 7.2 | 8.4 |
HeadL | 22.1 | 24.4 | 24.4 | 25.3 |
HeadW | 14.5 | 16.0 | 17.7 | 16.4 |
HeadD | 9.5 | 9.2 | 10.3 | 10.7 |
RosW | 3.8 | 3.7 | 3.6 | 3.9 |
RosH | 2.0 | 1.5 | 1.9 | 1.9 |
MenW | 2.9 | 2.9 | 3.2 | 3.1 |
MenL | 1.8 | 1.8 | 2.3 | 2.9 |
InterN | 2.3 | 2.8 | 2.5 | 2.7 |
SnOrb | 9.1 | 9.5 | 9.5 | 9.6 |
NosOrb | 6.3 | 6.7 | 6.3 | 7.0 |
OrbD | 7.7 | 7.5 | 8.2 | 7.2 |
InterOrb | 3.8 | 3.9 | 4.2 | 4.1 |
OrbEar | 6.2 | 6.9 | 7.6 | 7.1 |
EarL | 0.8 | 0.8 | 1.3 | 0.9 |
NeckW | 6.7 | 6.8 | 8.7 | 9.7 |
FaL | 10.9 | 11.3 | 11.0 | 10.8 |
TibL | 15.4 | 14.6 | 15.3 | 15.8 |
DigitL I–V | 4.4–5.9–6.6–6.8–6.1 | 4.3–NA–5.8–6.2–6.0 | 5.6–6.1–7.8–7.8–6.4 | 4.8–6.3–7.2–7.2–6.5 |
Relative DigitL | IV> III> V> II> I | NA | IV = III> V> II> I | IV = III> V> II> I |
ToeL I–V | 4.0–6.5–7.2–7.6–7.5 | 4.0–6.3–6.9–7.7–7.3 | 5.1–7.1–7.4–8.4–7.7 | 5.3–6.4–6.3–8.4–8.2 |
Relative ToeL | IV> V> III> II> I | IV> V> III> II> I | IV> V> III> II> I | IV> V> II> III> I |
SL | 10/11 | 11/10 | 10/12 | 12/11 |
SLMOrb | 8/7 | 8/8 | 8/8 | 9/8 |
IL | 11/10 | 10/11 | 9/10 | 9/10 |
InterCilS | 37 | 37 | 37 | 35 |
InterOrbS | 18 | 17 | 15 | 18 |
DigitLa I–V (left manus) | (5+10)–(4+10)–(4+12)– (5+12)–(6+11) | (4+8)–(4+10)–(4+11)– (5+11)–(4+10) | (4+9)–(4+10)–(4+12)– (5+10)–(5+9) | (4+8)–(4+9)–(4+12)– (4+10)–(4+9) |
DigitLa I–V (right manus) | (5+10)–(4+12)–(4+12)– (5+10)–(6+11) | (3+7)–(4+9)–(4+13)– (5+11)–(4+11) | (4+7)–(4+8)–(4+11)– (5+9)–(5+9) | (4+7)–(4+7)–(4+10)– (4+11)–(4+9) |
ToeLa I–V (left pes) | (3+10)–(4+11)–(5+14)– (6+13)–(5+15) | (4+9)–(4+9)–(4+10)– (6+10)–(5+9) | (3+9)–(4+10)–(5+11)– (6+12)–(4+12) | (3+8)–(4+9)–(4+12)– (6+11)–(5+12) |
ToeLa I–V (right pes) | (3+10)–(4+11)–(5+13)– (6+12)–(6+15) | (5+10)–(4+11)–(4+14)– (5+13)–(5+14) | (3+8)–(4+10)–(4+12)– (6+11)–(5+12) | (3+9)–(3+8)–(3+12)– (5+12)–(5+12) |
DTR | 18 | 18 | 18 | 18 |
PV | 30 | 32 | 33 | 29 |
PV’ | 23 | 24 | 25 | 22 |
VentR | 42 | 40 | 42 | 40 |
FemPreEnSc | 13+13 (diastema 1 scale) | 14+14 (diastema 1 scale) | 34 (no diastema) | 32 (no diastema) |
FemPre Po/Pi | 2 PrePo+2 PrePo | 2 PrePo+2 PrePo | 6 PrePi | 6 PrePi |
Postcloacal spurs | 2/2 | 3/2 | 2/2 | 3/2 |
characters are given left/right; paired measurements are given for the right side. NA = not assessed / not available.
No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.
Kingdom |
|
Phylum |
|
Class |
|
Order |
|
Family |
|
Genus |
Cyrtodactylus panitvongi
Pauwels, Olivier S. G., Chotjuckdikul, Natthaphat, Donbundit, Nattasuda, Sumontha, Montri & Meesook, Worawitoo 2024 |
Cyrtodactylus panitvongi
Pauwels & Chotjuckdikul & Donbundit & Sumontha & Meesook 2024 |
Cyrtodactylus panitvongi
Pauwels & Chotjuckdikul & Donbundit & Sumontha & Meesook 2024 |
Cyrtodactylus panitvongi
Pauwels & Chotjuckdikul & Donbundit & Sumontha & Meesook 2024 |
Cyrtodactylus panitvongi
Pauwels & Chotjuckdikul & Donbundit & Sumontha & Meesook 2024 |
Cyrtodactylus panitvongi
Pauwels & Chotjuckdikul & Donbundit & Sumontha & Meesook 2024 |
Cyrtodactylus panitvongi
Pauwels & Chotjuckdikul & Donbundit & Sumontha & Meesook 2024 |
Cyrtodactylus panitvongi
Pauwels & Chotjuckdikul & Donbundit & Sumontha & Meesook 2024 |
C. denticulatus
Chomdej, Suwannapoom, Pradit, Phupanbai & Grismer 2023 |
C. thongphaphumensis
Grismer, Rujirawan, Chomdej, Suwannapoom, Yodthong, Aksornneam & Aowphol 2023 |
C. fluvicavus
Grismer, Aowphol, Yodthong, Ampai, Termprayoon, Aksornneam & Rujirawan 2022 |
C. rivularis
Grismer, Aowphol, Yodthong, Ampai, Termprayoon, Aksornneam & Rujirawan 2022 |
C. uthaiensis
Grismer, Aowphol, Yodthong, Ampai, Termprayoon, Aksornneam & Rujirawan 2022 |
C. rukhadeva
Grismer, Suwannapoom, Pawangkhanant, Nazarov, Yushchenko, Naiduangchan, Le, Luu & Poyarkov 2021 |
C. ngati
Le, Sitthivong, Tran, Grismer, Nguyen, Le, Ziegler & Luu 2021 |
C. muangfuangensis
Sitthivong, Luu, Ha, Nguyen, Le & Ziegler 2019 |
C. muangfuangensis
Sitthivong, Luu, Ha, Nguyen, Le & Ziegler 2019 |
C. samroiyot
Pauwels & Sumontha 2014 |
C. kunyai
Pauwels, Sumontha, Keeratikiat & Phanamphon 2014 |
C. kunyai
Pauwels, Sumontha, Keeratikiat & Phanamphon 2014 |
C. auribalteatus
Sumontha, Panitvong & Deein 2010 |
C. tigroides
Bauer, Sumontha & Pauwels 2003 |
C. chanhomeae
Bauer, Sumontha & Pauwels 2003 |
C. interdigitalis
Ulber 1993 |