Copy igh © Ame ican Museum o Na u al His o y 2019 ISSN 0003-0082
AMERICAN MUSEUM NOVITATES
Numbe 3929, 14 pp. June 19, 2019
Desc ip ion o a New Species o Pony ish
(Teleos ei: Leiogna hidae: Equuli ini: Pho ola e alis)
om he Gul o Oman
JOHN S. SPARKS1 AND PROSANTA CHAKRABARTY2
ABSTRACT
A new species belonging o he leiogna hid genus Pho ola e alis, collec ed om he coas al
wa e s o Oman, is desc ibed he ein. Pho ola e alis is unique among leiogna hid gene a in pos-
sessing a species-speci ic anslucen midla e al lank s ipe ha may be comp ised o ei he
mul iple independen anslucen windows (P. s e co a ius, P. mo e oniensis, and he new spe-
cies) o a con inuous anslucen la e al band (P. an ongil). Pho ola e alis poly enes us, new
species, is dis inguished om congene s by he p esence o a sho , composi e midla e al s ipe
comp ised o h ee small, ounded anslucen windows ( s. nume ous windows in bo h P.
s e co a ius and P. mo e oniensis, o a con inuous anslucen s ipe in P. an ongil), and ha is
con ined o he mid lank ( s. ex ending a majo i y o he leng h o he lank in congene s). The
new species is u he dis inguished om bo h P. mo e oniensis and P. an ongil by a shallowe
body, and om bo h P. s e co a ius and P. mo e oniensis by a pigmen a ion pa e n abo e he
la e al midline comp ised p ima ily o la ge ounded blo ches ( s. smalle sinuous lines and
ma kings comp ising a e micula ed pa e n). Pho ola e alis poly enes us is cha ac e ized by a
lowe jaw ha is deep and con ex in la e al iew ( s. mos ly s aigh in congene s, excluding P.
an ongil), and ha o ms an angle o be ween 60°–70° o ho izon al ( s. less han 45° in con-
gene s, excluding P. an ongil). The only o he species o Pho ola e alis epo ed om he Wes -
e n Indian Ocean is P. an ongil, o da e only known om he coas al wa e s o Madagasca ,
whe eas bo h P. s e co a ius and P. mo e oniensis ha e anges es ic ed o he wes e n Paci ic
and ex ending in o he eas e n Indian Ocean.
1 Depa men o Ich hyology, Di ision o Ve eb a e Zoology, Ame ican Museum o Na u al His o y.
2 Museum o Na u al Science (Fish Sec ion), Depa men o Biological Sciences, Louisiana S a e Uni e si y.
2 AMERICAN MUSEUM NOVITATES NO. 3929
INTRODUCTION
The genus Pho ola e alis was ecen ly desc ibed by Spa ks and Chak aba y (2015) o
encompass membe s o Leiogna hidae ha a e cha ac e ized by a species-speci ic anslucen
midla e al lank s ipe. Pho ola e alis cu en ly comp ises h ee species, P. s e co a ius (E e -
mann and Seale, 1907), P. mo e oniensis (Ogilby, 1912), and P. an ongil (Spa ks, 2006), no
including he new species desc ibed he ein. This cha ac e is ic lank s ipe may ei he be com-
p ised o mul iple independen anslucen windows (P. s e co a ius, P. mo e oniensis, and he
new species) o is a con inuous anslucen la e al band (P. an ongil), h ough which bac e ially
media ed luminescence is p esumably ansmi ed om he ligh o gan ia clea ing o he
sil e y la e al lining o he gas bladde (Spa ks e al., 2005; Spa ks, 2006; Spa ks and Chak ab-
a y, 2015). All membe s o Pho ola e alis we e o me ly placed in Equuli es (Kimu a e al.,
2008; Chak aba y e al., 2011a, 2011b), and Pho ola e alis sha es he p esence o anslucen
la e al lank egions wi h i s sis e genus Equuli es, al hough he mo phology o hese anslu-
cen egions di e subs an ially be ween he wo gene a. Pho ola e alis is eco e ed as he sis e
g oup o Equuli es, and oge he hese wo gene a comp ise he ibe Equuli ini (Chak aba y
e al., 2011a, 2011b), which is he sis e axon o he ibe Nuchequulini wi hin he sub amily
Gazzinae (Chak aba y and Spa ks, 2015).
Membe s o Pho ola e alis can be dis inguished om hei sis e axon Equuli es, as well as
o he membe s o he amily, by bo h ex e nal and in e nal ea u es o hei ligh -o gan sys em.
Ex e nally, male membe s o Equuli es a e cha ac e ized by an expansi e and anslucen i-
angula , co nucopia-shaped, o apezoidal pa ch on he lank (Spa ks and Chak aba y, 2007:
ig. 1), e sus a con inuous o composi e midla e al s ipe comp ised o mul iple independen
anslucen windows in Pho ola e alis (Spa ks and Chak aba y, 2015: ig. 3). In Pho ola e alis,
he anslucen la e al s ipe is ei he lacking en i ely in emales, o is conside ably less de el-
oped (no e: only he holo ype, a male, is known o he new species). Bo h Equuli es and Pho-
ola e alis exhibi la e al clea ing (= anslucence) o he guanine-lined in e nal gas bladde
su ace, and he ex en o clea ing co esponds o he loca ion o he anslucen ex e nal
pa ches o s ipe. La e al gas bladde clea ing in Equuli es, howe e , is a mo e ex ensi e,
ex ending he ull leng h o he gas bladde , whe eas in Pho ola e alis clea ing is es ic ed o
he pos e io egion o he gas bladde (see Spa ks e al., 2005: igs. 4, 5; Spa ks and Chak a-
ba y, 2015: igs. 2, 4). Male membe s o Pho ola e alis exhibi a mode a ely enla ged donu -
shaped ligh o gan ha ex ends a mos only sligh ly pos e io ly in o he gas bladde space,
whe eas he ligh o gan in male membe s o Equuli es is g ea ly enla ged, wi h pai ed do so-
la e al lobes ex ending pos e io ly well in o he gas bladde space (less E. leuciscus) (see Spa ks
and Chak aba y, 2015: igs. 2, 4).
Based on a numbe o axonomically comp ehensi e amily-le el phylogene ic hypo heses
ha ha e been gene a ed in ecen yea s, we now ha e a be e unde s anding o leiogna hid
di e si y, in a amilial ela ionships, and he e olu ion o hei unique ligh -o gan sys em
(Spa ks and Dunlap, 2004; Spa ks e al., 2005; Chak aba y e al., 2011a, 2011b). As a esul o
hese phylogene ic s udies, ex ensi e ieldwo k, and de ailed mo phological compa isons, se -
e al new leiogna hid gene a and species ha e ecen ly been desc ibed (e.g., see Chak aba y
2019 SPARKS AND CHAKRABARTY: NEW SPECIES OF PONYFISH 3
and Spa ks, 2008; Spa ks and Chak aba y, 2015; Suzuki and Kimu a, 2017). In he p esen
s udy, a new species belonging o Pho ola e alis is desc ibed om ma e ial collec ed in he Gul
o Oman.
MATERIALS AND METHODS
Mo phome ic measu emen s we e eco ded o he nea es 0.1 mm using dial calipe s.
S anda d leng h (SL) is used h oughou . Ve eb al coun s exclude he u al cen um (= las
hal -cen um). The i s caudal e eb a is he e de ined as he i s e eb a bea ing a de ini e
hemal spine (Hubbs and Lagle , 1949). Ve eb al and in-spine/ ay coun s we e ob ained om
adiog aphs. The e minal do sal- in and anal- in ays, which a e b anched o he base o he
in, a e coun ed as a single elemen .
Mo phology o he ligh -o gan sys em was examined in si u (new species) o in isola ion
ia dissec ion o pe mi de ailed ana omical compa isons. Sex o he holo ype was con i med
ia dissec ion. Os eological and so issue ea u es we e analyzed using s anda d ilm and digi-
al adiog aphs, high- esolu ion digi al images, MRIs (Chak aba y e al., 2011a), examina ion
o whole alcohol-p ese ed specimens, specimens clea ed-and-s ained (C&S) o bone and
ca ilage ( ollowing he p o ocol o Taylo and Van Dyke, 1985), and d y skele al (S) p epa a-
ions. Specimens examined a e lis ed unde Ma e ial Examined in appendix 1. Ins i u ional
abb e ia ions a e as lis ed in Le i on e al. (1985) and Sabaj Pé ez (2014).
RESULTS
Sys ema ic Accoun
Pho ola e alis poly enes us, new species
Figu es 1, 2; able 1
Holo ype: AMNH 267152, 66.8 mm SL, male; Sul ana e o Oman: Gul o Oman, Fish
Ma ke , 23° 36′ N; 058° 33′ E; Z. Baldwin, July 2008.
Diagnosis: Pho ola e alis poly enes us is dis inguished om congene s by he p esence o
a sho , composi e midla e al s ipe comp ised o h ee small, ounded anslucen windows
ha a e peppe ed wi h ch oma opho es ( s. nume ous windows in bo h P. s e co a ius and P.
mo e oniensis, o a con inuous anslucen s ipe in P. an ongil) and is con ined o he mid lank
( s. ex ending mos o en i e he leng h o he lank in congene s). The new species is u he
dis inguished om bo h P. s e co a ius and P. mo e oniensis by a pigmen a ion pa e n abo e
he la e al midline comp ised o la ge ounded blo ches and ma kings ( s. smalle sinuous
lines and ma kings o ming a e micula e pa e n), and bo h P. mo e oniensis and P. an ongil
by a shallowe body. Pho ola e alis poly enes us exhibi s a lowe jaw ha is deep and con ex
in la e al iew ( s. mos ly s aigh in congene s, less P. an ongil), and ha o ms an angle o
be ween 60°–70° o ho izon al ( s. less han 45° in congene s, less P. an ongil). The only o he
species o Pho ola e alis epo ed om he wes e n Indian Ocean is P. an ongil, o da e only
known om he coas al wa e s o Madagasca , and whe eas bo h P. s e co a ius and P. mo e oni-
4 AMERICAN MUSEUM NOVITATES NO. 3929
ensis ha e mo e ex ensi e anges, ne e heless hey a e es ic ed o he wes e n Paci ic and
ex ending in o he eas e n Indian Ocean.
Desc ip ion: P opo ional measu emen s and me is ic da a p esen ed in able 1 and in e -
nal os eological ea u es can be isualized in igu e 2. A mode a ely shallow-bodied and elon-
ga e leiogna hid. Body la e ally comp essed. La e al snou ou line s aigh wi h a p o ube ance
abo e uppe lip. Mou h small and e minal in posi ion. P emaxilla long and p o uding along
do sal ma gin. Pos e io ma gin o maxilla exposed, eaching o le el o e ical h ough an e-
io ma gin o o bi . Lowe jaw deep and no iceably con ex in la e al iew, o ming angle o
60°–70° o ho izon al. Mild p eo bi al p o ube ance due o p o usion o bo h on al and
la e al e hmoid ossi ica ions. P edo sal head p o ile weakly sloping and s aigh . Nuchal spine
(exposed ising do sal aspec o he sup aoccipi al c es ) e y sligh ly p o uding and dis al ip
exposed. Nuchal spine wi h dis inc median keel. Eye la ge. An e io nasal po e small and
ound; pos e io o amen much la ge and c escen shaped, pa ially enci cling an e io po e.
Two sho and s ou pos nasal spines p esen on la e al e hmoid, loca ed pos e io o nasal
o amina and jus os odo sal o o bi . Pos nasal spines ollowed pos e io ly by well-de eloped
sup ao bi al idges ha con e ge pos e io ly. One sup aneu al p esen wi h s ong an e io
p ocess. P eope cula ma gin smoo h, no se a e, along en al and en ocaudal ma gins.
Do sal and en al body p o iles mildly ounded, wi h en al p o ile mo e con ex han do sal.
Do sal- in o igin loca ed well pos e io o e ical h ough pel ic- in o igin. Anal- in o igin
loca ed a abou le el o e ical h ough he las (= 8 h) do sal- in spine/ i s do sal- in ay.
Caudal peduncle slende and shallow. To al e eb al coun 23: 10 p ecaudal + 13 caudal. Neu-
al and hemal spines o e eb al cen um PU4 weakly expanded dis ally and bladelike. Thi -
een elonga e and iangula ou e gill ake s a ayed along lowe limb (= 5 h ce a ob anchial)
o i s gill a ch.
Fins: Do sal in wi h VIII spines and 16 b anched ays. Fi s do sal- in spine g ea ly educed
in leng h and somewha obus . Second h ough ou h do sal- in spines elonga e and obus ;
second spine longes . Second do sal- in spine mode a ely, bu no exceedingly, elonga e com-
pa ed o hi d and ou h. Thi d and ou h do sal- in spines se a e along an e io ma gin
“lock” in o g oo e on pos e io ma gin o p eceding spine when e ec ( ig. 2). Do sal- in spines
i e h ough eigh eeble, conside ably sho e han second h ough ou h spines. Anal in wi h
III spines and 14 b anched ays. Fi s anal- in spine e y sho . Second and hi d anal- in spines
obus and elonga e. Thi d anal- in spine se a e on an e io ma gin “locks” in o g oo e on
pos e io ma gin o second spine when e ec ( ig. 2). Spinous do sal and anal ins wi h asqua-
ma e basal shea h. Eigh een o al pec o al- in ays. Pel ic ins sho , no eaching i s anal- in
spine when adduc ed (i.e., a signi ican gap p esen ). Eigh uppe and se en lowe b anched
caudal- in ays.
Den i ion: Two dis inc ows o ex emely closely se , elonga e and ecu ed conical ee h
p esen in bo h uppe and lowe jaws. Uppe jaw ee h somewha la ge and mo e ecu ed
han hose o lowe jaw. Lips leshy, and ee h embedded.
Squama ion: Body scales cycloid. Head and ope cula egion asquama e. Ches ully scaled,
excep along en al midline. Scales conspicuous. Squama ion ex ends an e io ly, en al o
2019 SPARKS AND CHAKRABARTY: NEW SPECIES OF PONYFISH 5
TABLE 1. Mo phome ic and Me is ic Da a o Holo ype o Pho ola e alis poly enes us.
Cha ac e Holo ype
S anda d leng h (mm) 66.8
Pe cen age o SL
Head wid h (max.) 13.2
Head leng h 27.7
Body dep h 35.9
Caudal peduncle leng h 5.4
Caudal peduncle wid h 2.3
Caudal peduncle dep h 5.8
Pec o al- in leng h 19.5
P edo sal leng h 41.5
P eanal leng h 53.4
P epel ic leng h 33.1
Pe cen age o HL
Snou leng h 33
O bi diame e 34.6
Uppe jaw leng h 22.7
Lowe jaw leng h 58.4
In e o bi al wid h 30.8
Caudal peduncle leng h/wid h 2.4
Caudal peduncle leng h/dep h 0.9
Gill ake s (lowe limb 1s a ch) 13
Ve eb ae (p ecaudal + caudal) 10 + 13 = 23
Do sal in VIII 16
Anal in III 14
ope cle, o app oxima ely an e io ma gin o ches . La e al line a ched and comple e, loca ed
well do sal o la e al midline. Po ed scales in la e al line highly embedded and di icul o
isualize and coun . Po es well de eloped. All ins asquama e, excep o caudal in, which bea s
se e al ows o cycloid scales o educed size, pa icula ly cen ally on in.
Pigmen a ion in P ese a i e: Body g ound colo a ion oli e o ligh b own abo e la -
e al midline, and sil e y, guanine co e ed below midline ( ig. 1). Pigmen a ion pa e n abo e
la e al midline cha ac e is ically blo chy and mo led, comp ised o da ke b own oundish
blo ches and b oad lines o e oli e o ligh b own backg ound. Blo ches a iable in size and
somewha closely a anged, pa icula ly os odo sally. Body jus en al o la e al midline
peppe ed wi h melanopho es. Ope cle, in e ope cle, and lac imal i idescen sil e y wi h golden
hue. P eope cle, subope cle, and cheek i idescen and sil e y. Sho , composi e, midla e al
s ipe p esen , comp ised o h ee oblong da k g ay anslucen windows s udded wi h ch o-
ma opho es, and appea ing much da ke han su ounding egion due o concen a ion o
6 AMERICAN MUSEUM NOVITATES NO. 3929
FIG. 1. Pho ola e alis poly enes us, new species, holo ype, AMNH 267152, male, 66.8 mm SL; Sul ana e o
Oman: Gul o Oman, Mu ah Fish Ma ke . No e: Specimen has ield ag on pos e io lank and an e io mos
midla e al window is obscu ed behind pec o al- in ays in image.
melanopho es. S ipe loca ed a mid lank, beginning abou le el o e minal adduc ed pel ic- in
ays. Lips g ayish. Lowe jaw and gula egion g ayish. Snou wi h la ge black pa ch abo e
uppe lip. Two dis inc black blo ches p esen an e io o o bi ; do sal blo ch co e ing an e io
po ion o pos nasal spines on la e al e hmoid. Do sal and en al ma gins o o bi lined in
black pigmen . In e o bi al egion and nape g ayish an e io ly o g ayish-b own pos e io ly.
La ge, oblong da k b own ma king pos e io o o bi and do sal o ope cle. Ches and belly
sil e y; sil e y along en al midline. Do sal and anal ins hyaline o pale yellowish-whi e, wi h
a sligh concen a ion o black pigmen dis ally on do sal- in spines. Base o do sal in black;
becoming mo e p ominen (i.e., da ke ) pos e io ly. Pec o al and pel ic ins yellowish an.
Caudal peduncle pale annish yellow wi h blackish saddle on do sal ma gin. Base o caudal in
pale yellow. Caudal in yellowish o oli e p oximal o base, becoming anslucen o whi e
dis ally. Memb ane o caudal in wi h concen a ion o black pigmen , c ea ing se ies o hin,
black adia ing lines, pa icula ly nea base.
Ligh -O gan Sys em (LOS): In Pho ola e alis he ligh -o gan sys em is in e nally cha -
ac e ized by a mode a ely enla ged, donu -shaped, and conspicuously spo ed ligh o gan in
males ha ex ends only sligh ly pos e io ly in o he gas bladde . In con as , in membe s o
Equuli es, i s sis e axon, he do sola e al lobes o he ligh o gan in males a e g ea ly enla ged,
hea ily pigmen ed, and ex end pos e io ly well in o he gas bladde . In e nally, male membe s
o Pho ola e alis exhibi pos e ola e al clea ing o he sil e y gas bladde lining, co esponding
in ex en o he ex e nal anslucen la e al s ipe o windows on he lank. The ligh -o gan
sys em in he new species is simila o ha desc ibed o males o o he membe s o Pho ola e-
alis (see Spa ks, 2006; Spa ks and Chak aba y, 2015). In he new species, la e al clea ing o
he gas bladde lining co esponds o he h ee ex e nal anslucen midla e al windows, he
an e io mos window loca ed a abou he midpoin o he adduc ed pec o al- in ays. Ex e -
nally, membe s o Pho ola e alis a e dis inguished om all o he leiogna hids by he p esence
o a anslucen lank s ipe, which, depending on he species, may be comp ised o ei he
2019 SPARKS AND CHAKRABARTY: NEW SPECIES OF PONYFISH 7
FIG. 2. Radiog aph o Pho ola e alis poly enes us, new species, holo ype, AMNH 267152, male, 66.8 mm SL;
Sul ana e o Oman: Gul o Oman, Mu ah Fish Ma ke .
mul iple independen anslucen windows (P. s e co a ius, P. mo e oniensis, and he new spe-
cies) ( igs. 1, 3A–C), o a con inuous midla e al band (P. an ongil) ( ig. 3D). In Pho ola e alis,
he anslucen la e al s ipe is ei he lacking en i ely in emales, o is conside ably less well
de eloped.
Dis ibu ion: Known only om a single specimen acqui ed a a ish ma ke (23° 36′ N;
058° 33′ E) in no heas e n Oman, Sul ana e o Oman. The new species is p esumably mo e
widesp ead in he egion as well as in he Gul o Oman. Ex ensi e ecen su eying and col-
lec ing in he adjacen po ion o he Pe sian (A abian) Gul did no esul in any addi ional
obse a ions o his species (pe sonal obs.).
E ymology: Named in e e ence o he cha ac e is ic anslucen midla e al windows, which
a e ounded and appea ing spo like. The speci ic epi he , poly enes us, is used as an adjec i e.
DISCUSSION AND COMPARISONS
As many as 10 leiogna hid species ha e been epo ed o occu in he coas al wa e s o
Oman (Randall, 1995; Al-Ju aili e al., 2010, F oese and Pauly, 2019). In addi ion, ecen s udies
ha e con i med he occu ence o bo h E. elonga us (Jayabalan e al., 2010; Jawad e al., 2013),
which had p e iously been epo ed om Oman, and E. klunzinge i (Jawad e al., 2012), new
eco d, om he coas al wa e s o Oman, Al hough bo h o hese species a e shallow-bodied,
elonga e pony ishes, hey a e cha ac e ized by an expansi e anslucen iangula o apezoi-
dal pa ch on he lank in males (Spa ks and Chak aba y, 2007: ig. 1a, d).
The only species Pho ola e alis poly enes us could po en ially be misiden i ied as and con-
used wi h (and ha has been epo ed o occu in he egion) is he species commonly e e ed
o as Leiogna hus oblongus o Equuli es oblongus (Valenciennes, 1835) ( ig. 4). No e, we use
Equuli es oblongus o con enience only, as he e a e no ex e nal apomo phic ea u es ha can
8 AMERICAN MUSEUM NOVITATES NO. 3929
FIG. 3. Species o Pho ola e alis illus a ing pheno ypic a iabili y o anslucen midla e al
lank s ipe cha ac e is ic o genus. (A) P. s e co a ius, USNM 55906, holo ype, 83.8 mm SL,
Philippines. (B) P. s e co a ius, SIO 96-96, non- ype, 98.2 mm SL, Philippines. (C) P. mo e o-
niensis, AMS I.22983001, non ype, 76.9 mm SL, Aus alia. (D) P. an ongil, AMNH 236544,
holo ype, 75.0 mm SL, Madagasca .
2019 SPARKS AND CHAKRABARTY: NEW SPECIES OF PONYFISH 9
FIG. 4. Equula oblonga, holo ype, MNHN A-6754, 61.8 mm SL; Timo , Indonesia.
be iden i ied on he holo ype o place he species in Equuli es. The holo ype o E. oblongus is
in ac , nondissec ed, and lacking any e idence o anslucen lank pa ch(es) o a s ipe ha
would allow one o place i in he genus Equuli es. Based on o e all body shape, including
la e al p o ile o he lowe jaw and emaining lank pigmen a ion pa e n abo e he la e al
midline in p ese a ion, we belie e he holo ype o E. oblongus (MNHN A-6754) should ac u-
ally be placed in Pho ola e alis.
We u he no e, howe e , ha specimens a ibu ed o E. oblongus ( om Oman) a e much
deepe bodied han he holo ype o E. oblongus, exhibi a e y di e en pigmen a ion pa e n
on he lank abo e he la e al midline, and lack he anslucen la e al windows o s ipe ha
cha ac e ize he new species, and membe s o Pho ola e alis in gene al (compa e igs. 1 and 3
wi h ig. 4; also see Randall, 1995: 193, ig. 486). Based on ex e nal mo phology, he specimen
shown by Randall and iden i ied as E. oblongus (1995: ig. 486) appea s o be a emale E. leu-
ciscus. Equuli es leuciscus is a widesp ead species in he Indo-Wes Paci ic, wi h a ange epo -
edly ex ending om Eas A ica and Madagasca , o no he n Aus alia and New Caledonia,
o Vie nam, Taiwan, and China (pe sonal obs.); F oese and Pauly, 2019), bu ha has no p e i-
ously been epo ed om Oman.
The o iginal desc ip ion o E. oblongus is insu icien o dis inguishing his species om
o he membe s o Equuli ini, despi e he holo ype (MNHN A-6754) being in ela i ely good
condi ion ( ig. 4). In addi ion o he anslucen la e al lank windows p esen in he new spe-
cies, P. poly enes us can easily be dis inguished om he holo ype o E. oblongus by lank
pigmen a ion pa e n abo e he la e al midline ( ew la ge, oundish blo ches in E. oblongus s.
a mo e blo chy and a iable pa e n in he new species, comp ised o nume ous smalle and
mo e i egula da k, oundish ma kings), a deepe body, a s aigh p edo sal p o ile ( s.
ounded in E. oblongus), and a much deepe and conca e ( s. ela i ely s aigh and sho in
E. oblongus) la e al lowe jaw p o ile ( igs. 1–2, 4).
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o ia s anda d mail om:
Ame ican Museum o Na u al His o y—Scien i ic Publica ions
Cen al Pa k Wes a 79 h S ee
New Yo k, NY 10024
This pape mee s he equi emen s o ANSI/NISO Z39.48-1992 (pe manence o pape ).
