Bijdragen tot de Dierkunde, 52 (2): 155-168 — 1982

New data on Metaloricaria paucidens

from French Guiana and Surinam

(Pisces, Siluriformes, )

by

I.J. H. Isbrücker & H. Nijssen

Institute of Taxonomie Zoology, University of Amsterdam,

P.O. Box 20125, 1000 HC Amsterdam, The Netherlands

Abstract minae Regan, 1904, and Lithogeneinae Gosline,

1947. A large number of previously unrecorded specimens of mailed

of the Metaloricaria 1975 Harttiinae Isbrücker, (Lori- The former subfamilies Boeseman,

cariinae, tribe Harttiini, subtribe Metaloricariina) is com- 1971, and Acestridiinae Isbriicker & Nijssen, pared with the known specimens from French Guiana and 1974, are considered as tribes of the . Surinam. This results in the recognition of two subspecies,

viz., M. paucidens paucidens Isbrücker, 1975, from the rivers The Astroblepidae Bleeker, 1862 (■= Argeini

Oyapock and Maroni (French Guiana, Surinam), and M. p. Bleeker, 1862, and Cyclopidae Eigenmann, 1910) nijsseni (Boeseman, 1976) from the rivers Suriname, Sara- and & — both Nickerie and These Scoloplacidae Bailey Baskin, 1976 macca, Corantijn (Surinam). subspecies

ranked — were originally described as two different , even previously as subfamilies are excluded

within two different Metaloricaria paucidens Isbrü- genera: from the family Loricariidae (Isbriicker, 1980: cker, 1975, and Harttia nijsseni Boeseman, 1976. Morpho- 5 and 130). metric and meristic data, together with illustrations are given into of the two subspecies. Diagnostic characters of Metaloricaria The subfamily Loricariinae is subdivided

and within the Harttiini is in- are provided, its position four tribes: dicated. Loricariini Bonaparte, 1831,

Farlowellini 1958, Résumé Fowler,

Harttiini Boeseman, 1971, Une comparaison a été réalisée entre les exemplaires de pois- Acestridiini Isbriicker & Nijssen, 1974. sons-chats cuirassés du genre Metaloricaria Isbrücker, 1975 (Loricariinae, tribu Harttiini, subtribu Metaloricariina) déjà Harttiini usually have 12 branched caudal fin

connus française et du et de nom- (de Guyane Surinam), whereas the of rays (rarely 11), eighteen genera breux étude exemplaires pas signalés auparavant. Cette the Loricariini have 10 branched caudal conduit à distinguer deux sous-espèces, à savoir: M. pauci- always

dens des paucidens Isbrücker, 1975, rivières Oyapock et fin rays. Maroni française, Surinam) et M. nijsseni (Guyane p. The tribe Harttiini is subdivided into two sub- (Boeseman, 1976), des rivières Suriname, Saramacca, Nickerie tribes with the following genera: et Corantijn (Surinam). Ces sous-espèces avaient été précé-

demment décrites comme espèces distinctes, et même comme (1) H a r 11 i i n a Boeseman, 1971. appartenant à des genres distincts: Metaloricaria paucidens

Isbrücker, et Harttia Boeseman, The Harttia 1975 nijsseni 1976. On (a) genus-group (a depressed des données présente morphométriques et méristiques pour body; snout tip naked) comprises: les deux illustration. sous-espèces, en même temps qu’une Des Harttia Steindachner, 1876, caractères diagnostiques du genre Metaloricaria sont fournis,

Harttiella et on indique sa position dans le cadre des Harttiini. Boeseman, 1971,

Cteniloricaria Isbriicker & Nijssen, in

Isbriicker, 1979. INTRODUCTION The Sturi (b) s oma genus-group (a com-

The family Loricariidae Bonaparte, 1831, is pressed body; snout tip not naked )

presently subdivided into the subfamilies Lorica- comprises:

riinae Bonaparte, 1831, Hypostominae Kner, Sturisoma Swainson, 1838,

1853, Ancistrinae Kner, 1853, Hypoptopomatinae P. de Miranda Ribeiro,

Eigenmann & Eigenmann, 1890, Neoplecosto- 1939, - METALORICARIA PAUCIDENS 156 I. J. H. ISBRUCKER & H. NIJSSEN

Advancement of Tropical Research and the Pterosturisoma Isbrücker & Nijssen, (WOTRO) University of Amsterdam. 1978b, wish for the information, We to express our gratitude Isbrücker Sturisomatichthys & Nijssen, loan and/or gift of specimens, and for the hospitality

received during the of this to the following in Isbrücker, 1979. preparation study Bauchot and persons: Mrs. Dr. M. L. Mr. F. dAubenton i i Boeseman (2) M e t a 1 o r i c a r n a Isbrücker, 1979- (MNHN), Drs. M. J. P. van Oijen and Dr. M. and Mr. L. Wal- (RMNH), Mr. G. J. Howes (BMNH), Metaloricaria Isbrücker, 1975. 2-4 by Mr. schaerts (IRScNB). Figs. were photographed

drawn Mr. L. A. van der Laan (ZMA) and fig. 5 was by

Metaloricaria was described from 14 specimens J. Zaagman (ZMA).

of M. paucidens (type-species) by Isbrücker

(1975). This species is recently recorded from Metaloricaria Isbrücker, 1975

French Guiana by Boeseman (1982, pertaining Metaloricaria Isbriicker, 1975: 2 (original diagnosis; type-

to 2 described as Metalori- specimens). Thirty specimens species, by original designation and monotypy, Isbriicker, 1975). Harttia nijsseni by Boeseman (1976) were sub- caria paucidens

sequently assigned to Metaloricaria (Isbrücker & This is the of the sub- genus only representative Nijssen, 1978a). tribe Metaloricariina. Sixty-six previously unrecorded specimens from the of the subtribe Hart- Compared to genera 16 localities in French Guiana and Surinam were tiina (cf. Introduction), Metaloricaria has much available for this study. All the 112 specimens of shorter and more solid teeth, whereas the teeth Metaloricaria now known have been reviewed or Meta- are less bent near the crown. Moreover, examined. The of this is to purpose paper present with loricaria has bilobate crowns a small outer further descriptive and distributional data. As the inner the lobe and a large lobe, outer lobe originat- result of our comparison we recognize a single ing much lower along the tooth than in the Hart- with in the Meta- species two subspecies genus: tiina which possess a strongly bifurcate crown on loricaria p. paucidens from the river Oyapock each tooth, both crown lobes being about equally (= Oiapoque), bordering French Guiana and large (Isbriicker, 1975: 8, fig. 4). , Território do Amapâ, and from the river maximum teeth Metaloricaria has a of 26 (Boe- Maroni ( =Marowijne), bordering French Guiana table no. seman, 1976: 172, 5, specimen 28, and Surinam; the second subspecies is M. p. nijs- records 27 mandibular teeth in the holotype of seni which occurs in the rivers Suriname, Sara- the Harttia nijsseni, which we re-examined); macca, Nickerie and Corantijn in Surinam (the number of teeth increases with size, from 4 (in latter river is the boundary between Surinam and specimens of about 57 mm in SL) to 26 (in Guyana ). about in The specimens of 270-295 mm SL). Notes the and and illustra- on genus species, of the subtribe Harttiina have at least genera tions of both subspecies are included for direct 40 teeth in each jaw. The comparison. present publication is part of a the of the Compared to genera Harttiina, intended series to provide information for a revi- Metaloricaria is characterized by its premaxillae sion of the subfamily Loricariinae. The methods which do not meet at the symphysis, showing a and of taking measurements meristic data were furthermore it is distinct its large gap; by pre- defined by Isbriicker & Nijssen (1978a: 180-182). maxilla and dentary, which are much shorter in

transverse view. The material reported upon herein is deposited in the fol- In Metaloricaria is distinct its lowing institutions: British Museum (Natural History), Lon- addition, by don (BMNH), Institut Royal des Sciences Naturelles de papillose anterior and lateroventral side of the Brussels Belgique, (IRScNB), Instituut voor Taxonomische upper lip (fig. 1), which is conspicuous and Zoölogie (Zoölogisch Museum), Amsterdam (ZMA), Mu- broad in with the of the Hart- séum National d'Histoire Naturelle, Paris (MNHN), and comparison genera

Rijksmuseum van Natuurlijke Historie, Leyden (RMNH). tiina; its lower lip is longer and broader, whereas collected by The specimens Nijssen were assembled during the maxillary ( = rictal) barbel is distinctly longer the Biological Brokopondo Research Project (1966/1967), an in head in Metaloricaria expedition sponsored by the Netherlands Foundation for the (2.0-4.0 length against BIjDRAGEN TOT DE DIERKUNDE, 52 (2) - 1982 157

1. Metaloricaria anteroventral view of in ZMA Fig. paucidens nijsseni (Boeseman, 1976), a specimen 106.334, SL 255 mm, and of from showing the mouth lips which are characteristic the genus (reproduced Nijssen, 1970: 13).

in of sides and 5.0-10.7 Harttiina). The posterior margin outer of the premaxilla dentary ) a promi- the lower is convex concave at either is The lip medially, nently fleshy, papillose flap present. upper side towards the maxillary barbel. Upper and oral valve membrane is provided medially with lower lips, together with the maxillary barbels, an elongate membranaceous extension, which is form a horseshoe-like outline 1 in con- several times than that in ( fig. ), larger genera of the trast to the oval to roundish outline of the lips Harttiina. in the Harttiina; the lip shape of Metaloricaria is Apart from the differences mentioned above,

the Loricariidae. On the surface neither unique among morphometric characters, expressed as of the lip inside the buccal cavity (between the ratios, nor most of the counts distinguish Meta- 158 I. J. H. ISBRUCKER & H. NIJSSEN - METALORICARIA PAUCIDENS

of the Harttiina. The branches of the all loricaria clearly from genera subsidiary fin rays stem

main However, Metaloricaria comprises a larger species from one side of the (anterior) branch,

in than found in of the for the branch the (up to 295 mm SL) any except posterior in

related with which has a dichotomous distal genera. Cteniloricaria, specimens up rays, single

fin to 190 mm in SL contains the next to largest branching. Especially the third pectoral ray

species of the tribe. is broad (it has 9 subsidiary branches in the largest

The morphometric and meristic variation is specimen ).

summarized: The upper and next to upper branched caudal

fin thicker in rays are gradually distinctly larger

than in smaller this to a lesser SIZES: specimens; degree

with the lower and next to lower standard happens length 57.4-295.0 mm;

axial branched caudal fin well. length 63.7-322.2 mm; rays as

total 65.5-335.6 length mm; In smaller specimens the caudal fin is forked, smallest mature male 224.0 mm. than the lower lobe slightly or distinctly longer

the RATIOS OF STANDARD LENGTH: the upper lobe. The caudal fin, especially

is in head length 4.2-5.1; upper lobe, larger specimens frequently predorsal length 3.2-3.8; (more often than not) damaged during life, often postdorsal length 1.5-1.7; either rounded, obliquely truncate, or in various postanal length 1.8-2.0; the dorsal spine length 4.0-5.8; degrees of regeneration. The distal half of

dorsal length first fin 4.2-5.9; lobe has dark ray lower caudal fin often a prominent, anal spine length 5.8-8.8; brown blotch. Caudal triangular scutelets poste- pectoral fin spine length 4.6-6.4; with brown pelvic fin spine length 5.3-9.4; riorly a margin in smaller specimens;

length caudal "spine" 5.4-9.6; caudal upper fin often heavily spotted or with a few length lower caudal "spine" 5.6-10.2. ill-defined vertical bars in large specimens.

RATIOS OF HEAD LENGTH: No pectoral pore could be found.

snout length 1.6-2.0; Eye covered with a narrow dorsal fleshy length lower lip 3.1-6.2; (erroneously indicated as "skinny" by Isbriicker, thoracic length 1.5-1.8; 1975: 6) flap of skin, provided with minute, abdominal length 1.1-1.5; scattered odontodes maximum orbital diameter 3.7-5.8; in some specimens.

interorbital width 4.6-6.1; Additional characters were described by Is- cleithral width 1.0-1.2; briicker (1975) and by Boeseman (1976). supracleithral width 1.4-1.7;

head width 1.1-1.3;

head depth 2.1-2.9; Secondary sexual dimorphism (fig. body depth at dorsal fin origin 2.0-3.4;

— The 2). illustrated male shows more promi- body width at dorsal fin origin 1.3-1.8; body width at anal fin origin 1.4-2.1; nently developed odontodes along the side of caudal depth peduncle 9.0-18.7; and the the head on dorsum of the pectoral fin width caudal peduncle 6.1-17.7; and first 4 rays) than was known length maxillary barbel 2.0-4.0. (spine pre- viously (Isbriicker, 1975: 7 and 9, pis. I and III; COUNTS: Boeseman, 1976: 173, pl. 8). The pectoral fin lateral scutes 33-36; spines of males are thicker than those of females. coalescing lateral scutes 20-25; The largest odontodes the side of the head thoracic scutes 4-8; along premaxillary teeth 4-26; in the specimen, detail of which is illustrated in

mandibular teeth 6-26; 2 is about fig. 2.8 mm; the specimen figured in dorsal fin rays I, 6, i; 4 has in the same odontodes to about anal fin fig. region rays I, 4, i; up

fin 3.8 from pectoral rays I, 6; mm long; the odontodes arise mucous pelvic fin I, 5; rays skin and are difficult to measure accurately- caudal fin I (of 82 rays I, 12, specimens examined, 3 are In males short odontodes in ZMA enlarged occur on aberrant, 1 106.335 having I, 9, I, and 2 in ZMA & the ventral side fin in 106.335 106.338 having I, il, I caudal fin rays). of the pelvic spine, an BIJDRAGEN TOT DE DIERKUNDE, 52 (2) - 1982 159

2. Fig. Metaloricaria paucidens nijsseni (Boeseman, 1976), ￿ in ZMA 106.334, SL 256.1 mm, showing secondary sexual viz., odontodes the side of the head and dorsum dimorphism, enlarged along on the of the pectoral fin. 160 I. J. H. ISBRUCKER & H. NIJSSEN - METALORICARIA PAUCIDENS

other this erect position; they are very slender except for Compared to juveniles (see table II)

the thick, round tip, thus reminiscent in shape of smallest specimen examined has the following

fin and minute pins. The odontodes on the pelvic morphometric ratios meristic characters worth

in females. males anal fin spine are blunter than those In mentioning: spine 8.8; pelvic fin spine

the inner tooth lobe is somewhat shorter and its 9.4; length of lower lip 6.2; maximum orbital

than in females diameter fin distal tip is slightly more rounded 3.7; body depth at dorsal origin 3.4;

width fin width and juveniles. body at dorsal origin 1.8; body at

As indicated the smallest anal fin barbel already above, recog- origin 2.1; length maxillary 4.0;

male 224 in The 4 teeth in each mandibular teeth. nizable nuptial is mm SL. para- premaxilla; 6/7

Harttia type of nijsseni (ZMA 114.310, ex

listed Remark. — The and dentition of Meta- RMNH 27494, now: M. p. paucidens) was jaws

as specimen number 17 by Boeseman (1976: 172, loricaria are reminiscent of the structure usually

in of the table 5) and recorded with a SL of 223 mm. It present some members tribe Loricariini,

which in the subtribe Rineloricariina is a mature male, we measured with a SL viz., two genera of of 226 mm. Other mature male paratypes in Boe- Isbrücker, 1979 (Rineloricaria Bleeker, 1862,

seman's table 5 are the specimens number 22 and Isbrücker & Nijssen, in Is-

(RMNH 27502), 25 (RMNH 27496), 26 brücker, 1979), and in the subtribe Pseudolori-

(RMNH 27499), and 27 (RMNH 27495), with cariina Isbrücker, 1981.

recorded standard lengths between 239 and

265 mm. Discussion. — Metaloricaria was synon-

ymized with Harttia by Boeseman (1976: l68,

— M. He stated: "In Juvenile. In the juvenile topotype of p. 170). a previous paper (Boeseman,

in IRScNB there I omitted record of several paucidens 20029 (SL 57.4 mm), 1971), any specimens,

median the the Harttia-like is a ridge from tip of snout to be- clearly of a aspect, feeling some doubt

the anterior of the tween nostrils; a broad ridge runs to about their systematic allocation on account

the orbital rim, almost reaching the margin of relatively limited numbers of teeth, especially in

the snout. Dorsum of the head to about the Since I have come to the origin young examples. then, of the dorsal fin, and the sides of the caudal conclusion that these specimens, which evidently

Harttia peduncle comparatively more coarse than in larger represented a new species [ nijsseni, now

specimens. Abdomen naked except for a few considered Metaloricaria paucidens nijsseni],

scattered minute scutelets covered with should indeed be allocated the Harttia widely to genus

their the odontodes. The thoracic scutes (5/7 in number) on account of general shape, indistinct

of the and are very weakly developed. The tip snout rounded lateral longitudinal ridges, the

in adults is followed orbital the lack naked, ventroposteriorly by a missing notch, of any caudal

transverse series of well-developed scutelets. These filaments and, in adults, the still quite numerous

scutelets in are absent this juvenile. Supraorbital teeth found on each half jaw. The smaller number

raised. The in of teeth in in this margin conspicuously lips are as juveniles, not unexpected group,

smaller is adults except for the presence of relatively evidently of little systematic or phylogenetic and much less papillae. Papillae and fleshy pro- importance."

in the tuberances buccal cavity are exactly as in Boeseman (1976: 170) considered that the adults. Teeth with a very long oblong, distally distinguishing characters of Metaloricaria pauci-

rounded inner tip and with a minute, triangular dens (viz., the nature of the dentition and the

is acute outer tip. The specimen pale, with light shape and the structure of the lips) "...form a brown unit with pigment forming some indistinct spots on single functional presumably a con-

the anterior half of the and body, and some faint siderable adaptability therefore probably

transverse stripes on the dorsum to the sides of without sufficient importance to warrant generic

the caudal there is small dark Harttia peduncle; a spot on distinction from Steindachner. Especially

the caudal fin base. as many other diagnostic characters also con- BIJDRAGEN TOT DE DIERKUNDE, 52 (2) - 1982 161

vincingly show a close relationship of M. pauci- postorbital notch. The number of branched caudal

dens with Harttia surinamensis fin in 10 in my (Boeseman, rays (12 or 11 Harttiini, Loricariini)

is the characters to allocate 1971)." among most convincing

Even when we do not consider the dental and Metaloricaria.

Since Boeseman's of H. in labial characters of Metaloricaria, unique among description nijsseni

that Metaloricaria contained the Harttiini, we fail to understand Boeseman's 1976, we agreed two

solution this with Harttia. distinct As result of the to synonymize genus species. a present study,

Previously (1971) he omitted “Harttia” nijsseni we conclude that only a single species with two

his Harttiinae even from subfamily (or "comb- subspecies can be recognized.

whilst toothed" Loricariinae) at that time many

specimens were available to him. Boeseman ( 1971 : Metaloricaria paucidens Isbrücker, 1975 in 9, table 1) included this subfamily Harttiella,

Parasturisoma 1911" = A. de Harttia, "Ribciro, Discussion. — The earliest preserved speci-

Miranda Ribeiro, 1912 of Metaloricaria collected [a junior synonym men of paucidens was by

and as well as Farlowella Boeseman Boese- Sturisoma], Sturisoma, on December 28th, 1963 (cf.

latter is Eigenmann & Eigenmann, 1889 (the man, 1976: 171, in list of material). tribe presently assigned to the Farlowellini). Un- The first published record of Metaloricaria

like the of these “Harttia” is many species genera, paucidens an (at that time) unidentified photo-

is not "comb-toothed". — colour slide made in the nijsseni graph after a field

Boeseman two Suri- from — in previously (1971) assigned a freshly preserved specimen a popular

with much closer relation- nam species an evidently account of some Surinam freshwater fishes (Nijs-

into different Harttia surinamensis with ship genera: sen, 1970: 13), the caption in Dutch: "Zuig-

and Parasturisoma maculata Boeseman, 1971, bek van harnasmeerval" meaning "Suckmouth of

latter Boeseman, 1971 (the was subsequently mailed " (here reproduced in fig. 1).

into H. placed Cteniloricaria) . Both surinamensis For a review or for a more detailed examina-

and C. maculata less distinct from each other are tion, 112 specimens were at our disposal. On the

the (cf. Boeseman, 1971: 25, key to Surinam basis of previously published data (Isbriicker,

of than Meta- species Harttiinae) both are from 1975; Boeseman, 1976), which have been in-

loricaria Boeseman's table in paucidens. (1976: 173, corporated this paper, additional specimens

of “Harttia” “H.” 6) comparison paucidens, nijs- from different localities were taken at random for

and H. surinamensis is difficult seni, very to read, a complete examination. In the course of this,

the characters "U" and "V" although (number many characters were not taken from all of the

of scutes in series, and teeth number either because longitudinal specimens, of a considerable over-

on half upper/lower jaw, respectively; cf. his in ratios and counts or lap because of the great

fig. 1 on facilitate a clear distinction. differences in size page 154) (SL) of the specimens in the In a recent publication Boeseman various A however, (1982: samples. meaningful comparison be-

listed Metaloricaria 57) correctly paucidens from tween local populations of Loricariinae should be

a locality in French Guiana, without comments based the upon specimens of about same size.

its Most upon generic assignment. of our data are given in tables I-III. Several

Metaloricaria is reminiscent of Superficially, characters tend to differ locally, but we are unable

some member of the tribe Loricariini rather than to assess the value of these tendencies. For con-

of a member of the Harttiini. Like all venience of Harttiini, two size — comparison, groups one

a few of Loricariini lack a notch of species postorbital specimens over 150 mm in SL, the other of

Loricaria Von (e.g., piracicabae Ihering, 1907, under mm — are specimens 150 presented L. prolixa Isbrücker & Nijssen, and L.1978a, separately (in tables I and II, respectively), Isbrücker, cf. also lentiginosa 1979; Isbrücker, arranged according to the different river basins the 1981), although of the members of which Additional material majority they occupy. will be the Loricariini small possess a to quite prominent needed to fill the still existing gaps. 162 I. J. H. ISBRUCKER & H. NIJSSEN - METALORICARIA PAUCIDENS

Table I

Summary ofmorphometric and of some meristic characters of the two subspecies of Metaloricaria paucidens Isbrücker,

mm in river of 1975, specimens over 150 SL, arranged according to system. Data 7 specimens from the Oyapock, 7 from the Marowijne, 28 from the Suriname, 3 from the Saramacca, 5 from the Nickerie, 2 from the Sipaliwini, and from 3 the Corantijn river drainages are given, including data published previously. Of some of the specimens, selected characters Head lower caudal are given only. length through length “spine” are ratios of SL, snout length through

length maxillary barbel are ratios of head length.

M. M. subspecies p. paucidens paucidens nijsseni

drainage Oyapock Marowijne Suriname Saramacca Nickerie Sipaliwini Corantijn

Standard length (mm) 1158.5-237.0 171.0-270.0 155.0-295.0 231.0-248.0 164.0-266.2 257.0-274.0 180.0-203.0 head length 4.7-5.1 4.6-5.1 4.2-4.7 4.5-4.6 4.7-4.9 4.4-4.6 4.7-4.8 predorsal length 3.4-3.7 3.3-3.8 3.2-3.5 3.3-3.4 3.4-3.7 3.2-3.3 3.4-3.6

postdorsal length 1.5-1.6 1.5-1.6 1.6-1.7 1.6 1.6 1.7 1.6

postanal length 1.8-1.9 1.8-2.0 1.9-2.0 2.0 1.9 2.0 1.9

dorsal spine length 4.0-4.7 4.1-5.0 4.3-5.8 4.2-4.9 <6.0 4.3-5.0 4.4-4.6

first dorsal fin 4.3-4.7 4.2-4.9 4.7-5.7 4.6 5.9 5.3 length ray anal spine length 6.9-7.8 6.4-7.2 6.1-7.5 5.9-6.7 7.7 5.8-6.9 6.2-6.4

pectoral fin spine length 4.9-5.2 4.7-6.4 4.6-5.9 5.0-5.1 5.9 4.8-5.6 4.9-5.1 pelvic fin spine length 6.0-6.3 5.8-6.8 5.3-6.2 5.4-5.6 6.2 5.4-5.8 5.6-5.8

7.4-7.6 7.5 9.6 length upper caudal "spine" 5.4-<8.1 7.4-<9.1 <10.1 length lower caudal "spine" 6.0-7.4 5.7-8.1 6.7-<8.4 5.6 10.2 6.6

snout length 1.6-1.7 1.6-1.8 1.6-1.9 1.6-1.7 1.6 1.7-1.8 1.8

length lower lip 3.1-3.7 3.4-3.8 3.5-4.0 3.8 3.9 3.9 thoracic length 1.6-1.8 1.5-1.8 1.7-1.8 1.8 1.6 1.5 abdominal length 1.2-1.3 1.1-1.2 1.3-1.4 1.2 1.2 1.4 maximum orbital diameter 4.5-5.5 4.4-5.6 4.3-5.8 5.2-5.7 5.4 5.2-5.4 4.9-5.0

interorbital width 5.0-5.3 4.6-5.5 4.7-6.1 5.2-5.4 5.3 5.1-5.4 5.2-5.6

cleithral width 1.0-1.1 1.0-1.1 1.1-1.2 1.1 1.0 1.1

supracleithral width 1.4-1.5 1.4-1.5 1.5 1.5 1.4 1.4

head width 1.1-1.2 1.1-1.2 1.2 1.1-1.3 1.1 1.1-1.2 1.2

head depth 2.3-2.6 2.3-2.5 2.4-2.6 2.4 2.5 2.1-2.3 2.4

body depth at dorsal fin 2.1-2.5 2.1-2.5 2.2-2.6 2.2-2.4 2.4 2.0-2.1 2.2-2.4

body width at dorsal fin 1.3-1.4 1.3-1.4 1.3-1.4 1.3-1.4 1.3 1.4 1.4

body width at anal fin 1.5-1.6 1.4-1.5 1.5-1.6 1.4-1.5 1.5 1.5 1.5

depth caudal peduncle 12.7-14.2 9.0-13.8 11.4-14.2 11.7-12.8 13.0 13.5-13.7 13.5-14.0 width caudal peduncle 6.6-7.8 6.1-6.6 6.1-7.2 6.1 7.1 7.6

length maxillary barbel 2.0-2.5 2.4-2.5 premaxillary teeth 10-20 11-19 14-26 18-22 21-22 20-26 17 mandibular teeth 11-21 14-19 16-26 13-21 21-23 21-26 19-21

Of the two forms described originally as two tion, the two subspecies of Metaloricaria paucidens

M rather the different species ( . paucidens and H. nijsseni), are easily recognizable by following

the colour to be the easiest and differences in colour pattern appears pattern:

perhaps most reliable character to distinguish

— Dorsum and sides of and head with body many conspic- them, not provided that the specimens are faded dark brown broad bars uous spots; no transverse poste- in colour during The differences rior base of last dorsal fin about preservation. to ray (juveniles with be pattern proved to geographically correlated. three narrow transverse stripes on dorsum and sides of caudal peduncle) Moreover, the variability in details of the colour Metaloricaria p. paucidens Isbriicker, 1975 is pattern quite great. Therefore, we assume it is

— Dorsum and sides of body and head with more opportune and more realistic to consider many vague brown spots or without spots; often there are up to five paucidens and distinct rather nijsseni subspecies broad brown transverse or blackish bars present posterior than different base of last dorsal fin colour species. to ray (juveniles have a

pattern which is reminiscent much of that in juve- In addition to certain morphometrical (tables very niles of the nominate subspecies) I-II) and meristic (table III) geographical varia- Metaloricaria p. nijsseni (Boeseman, 1976) BIJDRAGEN TOT DE DIERKUNDE, 52 (2) - 1982 163

Table II

of Summary morphometric and of some meristic characters of the two subspecies of Metaloricariapaucidens Isbrücker,

under in river of 1975, specimens 150 mm SL, arranged according to system. Data 4 specimens from the Oyapock, 3 from the Marowijne, 4 from the Suriname, 2 from the Saramacca, 2 from the Nickerie, and 2 from the Corantijn

river are of drainages given, including data published previously. Presentation data is the same as in table I.

M. M. subspecies p. paucidens paucidens nijsseni

drainage Oyapock Marowijne Suriname Saramacca Nickerie Corantijn

Standard length (mm) 81.5-144.5 139.5-149.0 98.0-145.0 90.0-128.0 100.9-122.7 115.0-127.0

head length 4.5-5.1 5.0 4.7-4.9 4.7-4.8 4.7-4.8 4.9

predorsal length 3.5-3.7 3.7 3.5-3.6 3.4 3.6 3.6

postdorsal length 1.6 1.6 1.6 1.6 1.6 1.6

postanal length 1.8-1.9 1.8-1.9 1.9 1.9 1.9 1.9

dorsal spine length 4.7-4.9 4.7 4.2-4.7 4.5-4.6 4.6 4.1-4.4

first dorsal fin 4.8-5.2 4.4-5.1 5.0 length ray anal spine length 6.9-7.5 7.4-8.0 6.1-6.6 6.6-6.7 6.6 6.0-6.5 pectoral fin spine length 4.9-5.4 4.9-5.2 4.8-5.0 4.9 5.0 4.7-5.2

pelvic fin spine length 6.5-6.8 6.5-8.5 5.8-6.3 5.9-6.1 6.4 5.8-6.7

caudal 6.9-8.7 7.5-7.7 length upper "spine"

length lower caudal "spine" 6.3-7.9 6.3-7.7 <7.6

snout length 1.7-1.9 1.8 2.0 2.0 1.9 2.0

length lower lip 3.1-3.5 3.5-5.2 3.9 thoracic length 1.7 1.5-1.6 1.7

abdominal length 1.3-1.5 1.2-1.3 1.3

maximum orbital diameter 4.0-4.4 4.3-4.5 3.9-4.6 3.8-4.4 3.9 4.0-4.2

interorbital width 4.8-5.7 5.0-5.7 5.5-5.9 5.6-5.7 5.7 5.3

cleithral width 1.1-1.2 1.1 1.1

supracleithral width 1.5-1.7 1.5 1.5

head width 1.1-1.2 1.1 1.1-1.2 1.2 1.2 1.2

head depth 2.5-2.9 2.3-2.4 2.5-2.7 2.4-2.5 2.7 2.3-2.4

body depth at dorsal fin 2.5-2.8 2.2-2.4 2.4-2.8 2.6-2.8 2.9 2.5

body width at dorsal fin 1.5-1.6 1.3-1.4 1.3-1.5 1.4 1.4 1.4

body width at anal fin 1.6-1.9 1.5-1.6 1.5-1.7 1.5-1.6 1.6 1.5-1.6 depth caudal peduncle 14.3-18.7 11.5-13.5 13.9-16.9 16.1-17.5 15.4 15.7-17.0

width caudal peduncle 7.5-17.7 7.3-7.7 7.5

length maxillary barbel 2.2-2.9 2.4-3.2 2.4 premaxillary teeth 5-12 9-13 9-12 8-10 8-9 9-12

mandibular teeth 8-15 11-13 11-17 12-13 11 11-14

As noted below (under Metaloricaria p. nijs- Metaloricaria paucidens paucidens Isbrücker,

seni), specimens without broad transverse bars 1975

and even without prominent colour were tables pattern (Figs. 3, 5; I-III) encountered in Surinam. In the samples from the Metaloricaria paucidens Isbriicker, 1975: 2-9, figs. 4a-f, rivers and some Suriname, Saramacca, Corantijn I-III, table 1 13 pis. (original description; holotype, para- the of specimens (RMNH 27495, part of RMNH types; type-locality: "French Guiana, creek at right bank

of of = Ouaqui River, upstream Saut Bali, Maroni ( Maro- 27500, and part of RMNH 27494, respectively) river from the wijne, Surinam) system"; paratypes type- differ from other M. there- p. nijsseni (and are locality and from French Guiana, Marouini River and

fore identified River: Saut Alicoto and Sikini Creek; only tentatively as this subspecies) Oyapock com- with Harttia surinamensis and lack the parison Boeseman, 1971 by the of broad bands on caudal peduncle, Harttia maculata (Boeseman, 1971)); — Isbriicker & Nijs- and the faint but by possession of a narrow, well- sen, 1978a: 178 (discussion); — Isbriicker, 1979: 88

defined — — transverse line near the posterior margin (listed); Isbriicker, 1980: 96 (listed); Nijssen, Van ZMA each Tuijl & Isbriicker, 1982: 59 (paratype in 112.741 of middorsal body scute; these lines are listed); — Boeseman, 1982: 57 (listed; French Guyana, more gradually conspicuous posteriorly. Saut Gostou). 164 I. J. H. ISBRUCKER & H. NIJSSEN - METALORICARIA PAUCIDENS

Table III

Frequency of the number of lateral and coalescing scutes of the

two subspecies of Metaloricaria paucidens Isbrücker, 1975, arranged river The in according to system. specimens (number parentheses)

were counted on both sides. Data publishedpreviously are included;

of lateral Boeseman is to the number scutes given by (1976), one

added, because the count given here includes the small triangular

scutelet on the caudal fin base.

drainage lateral scutes coalescing scutes

33 34 35 36 20 21 22 23 24 25

M. p. paucidens

Oyapock (11) 1 11 7 3 0 0 10 8 3 1 Marowijne (10) 1 18 1 0 1 7 10 2 0 0

M. p. nijsseni

Suriname (32) 12 47 5 0 25 36 3 0 0 0 Saramacca (5) 1810 082000

Nickerie (2) 1300 022000 Sipaliwini (2) 0400 022000

Corantijn (5) 0640 226000

Harttia paucidens; Boeseman, 1976: 170-171, 173-174, table Remarks. — This subspecies notably differs

6 (discussion; with Harttia nijsseni comparison Boeseman, Metaloricaria of from p. nijsseni by the presence 1976 and Harttia surinamensis Boeseman, 1971; data after dark brown scattered on dorsum Isbrücker, 1975). profuse spots tables and sides of and head Harttia nijsseni; Boeseman, 1976: 170-174, 5-6 (in body (fig. 3); juveniles 4 from Surinam, Tapanahoni River, RMNH part; paratypes often have to three dark brown transverse up 52 27494); — Nijssen, Van Tuijl & Isbrüclcer, 1982: the caudal stripes on peduncle (cf. Isbriicker, (paratype in ZMA 114.310 [ex RMNH 27494] listed). 1975, pl. II).

Material examined. — In addition to the holotype Most of the morphometric and meristic data

(IRScNB 549), the 13 paratypes (IRScNB 550 through 553, are given in tables I-III. ZMA 112.741), and the 4 paratypes of Harttia nijsseni from The number of thoracic scutes in 11 Tapanahoni River, about 2 km below Paloemeu airstrip specimens

ZMA ex RMNH from the River basin is in (RMNH 27494, three; 114.310, one, Oyapock 6/4 one speci- examined the we have following specimens: 27494), 6 both sides in in men, on four, 6/7 or 7/6 four,

and both sides in In 7 on two specimens. 8 speci- FRENCH GUIANA: mens from the Marowijne River basin, there are IRScNB 20029 (topotype), SL 57.4 mm, Ouaqui River, right

bank tributary of the Tampok, Saut Bali, Maroni basin, coll. 6/7 or 7/6 thoracic scutes in four, 7 on both

J. P. Gosse, 18-XI-1969; — RMNH 28950, two, SL 125- sides in and both one, 7/8 or 8/7 in two, 8 on Falls, 206 mm, Gostou Maroni basin, coll. P. Planquette, sides in one specimen. 29-XI-1979 (recorded by Boeseman, 1982: 57).

MNHN SL 222 of 1982-733, mm, upper course Oyapock

River, Malipa itu, Yapakani, small creeks near Trois Sauts Metaloricaria (02°15'N, 52°53'W), coll. F. d'Aubenton, 28-IX-1976; — paucidens nijsseni (Boeseman,

ZMA SL 223 Saut 107.635, mm, Panacoupélou (02°43'N, 1976) 52°30'W), fall in upper Oyapock River, coll. F. d'Aubenton,

— tables 13-X-1976; MNHN 1982-734, SL 237 mm, Baton Pilon (Figs. 1, 2, 4, 5; I-III)

its confluence Creek, near (03°25'N, 52°08'W) with Harttia nijsseni Boeseman, 1976: 170-174, pl. 8, tables 5-6 Oyapock River, coll. F, d'Aubenton, 19-X-1976. (original description [in part]; holotype, 29 paratypes;

type-locality: "Sipaliwini River, south-eastern Surinam"; SURINAM: 1 paratype from the type-locality, paratypes from Surinam:

ZMA 106.341, SL 224 mm, district Marowijne, Maka Creek Awaradam (rapids), Gran Rio, upper Suriname River;

left bank of Lawa River, 10 km of of Stoelmans at S. centre Zandvallen, ca. 10 km above Avanavero Falls, Kabalebo

30-120 4 Broko- Island, depth cm, width m, running water, bottom River, Corantijn River basin; Suriname River near

coll. sand, H. Nijssen, 21-IV-1967. pondo, pools in dry (and in partly or mostly dry) river BIJDRAGEN TOT DE DIERKUNDE, 52 (2) - 1982 165

Metaloricaria in ZMA SL 234.5 from Saut Ali- Fig. 3. paucidens paucidens Isbrücker, 1975, paratype 112.741, mm (￿ coto, upstream of Camopi, Oyapock River basin), showing the colour pattern on the dorsum of the body and head, charac- teristic of the subspecies.

Metaloricaria ZMA SL 245 the colour Fig. 4. paucidens nijsseni (Boeseman, 1976), ￿ in 106.343, mm, showing pattern on the dorsum of the body and head, characteristic of the subspecies. & H. - METALORICARIA PAUCIDENS 166 I. J. H. ISBRUCKER NIJSSEN

below Data of the — Measurements in mm bed (and in shallow parts, subsiding water); pool holotype.

Feddiprati rapids, Saramacca River; Kwambaolo Creek, (followed between parentheses by the usual ratio): $, SL axial total length 62.8 right tributary of Sara Creek above Dam, Suriname River 274; length 301; 315.4; head length km 84.4 basin; [the paratypes from Tapanahoni River, ca. 2 (4.4); predorsal length (3.2); postdorsal length 160.9 dorsal fin downstream Paloemeu airstrip, are Metaloricaria p. pauci- (1.7); postanal length 135.5 (2.0); spine length

dens Isbriicker, 1975}; comparison with Harttia paucidens 54.4 (5.0); length first dorsal fin ray 51.9 (5.3); anal fin

and Harttia surinamensis). spine length 39.8 (6.9); pectoral fin spine length 48.9

Isbriicker fin 47.5 caudal fin Metaloricaria nijsseni; & Nijssen, 1978a: 178 (dis- (5.6); pelvic spine length (5.8); upper

88 lower caudal fin cussion); — Isbriicker, 1979: (listed); — Isbriicker, "spine" length >27.1 (<10.1); "spine"

41.4 37.6 lower 1980: 96 (listed). length (6.6); snout length (1.7); length lip 16.3 (3.9); thoracic length 41.3 (1.5); abdominal length

46.3 (1.4); maximum orbital diameter 12.1 (5.2); inter- Material examined. — In addition to the holotype orbital width 12.2 cleithral width 57.3 of (5.1); (1.1); supra- (RMNH 27498) and 25 the 29 paratypes (RMNH 27495 cleithral width 43.7 (1.4); head width 55.0 (1.1); head through 27497 and 27499 through 27504), we have examined depth 30.5 (2.1); body depth at origin of dorsal fin 31.4 the following specimens: (2.0); body width at origin of dorsal fin 46.1 (1.4); body

width at origin of anal fin 40.8 (1.5); depth caudal peduncle SURINAM: width caudal 4.6 (13.7); peduncle 8.3 (7.6); lateral scutes ZMA SL district Broko- 106.338, twenty, 75.3-169.5 mm, thoracic 34/34; coalescing scutes 22/22; scutes 6/6; pre-

Marowijne (= Gran) Creek, 63 km S. of dam at pondo, maxillary teeth 26/26; mandibular teeth 26/25 (Boeseman Afobaka, 150 bottom sand and depth cm, running water, recorded 27 mandibular teeth).

leaves, coll. H. Nijssen, 20-X-1966; — ZMA 106.344, two,

SL district Gran Mau 242-295 mm, largest a $, Brokopondo, Notes. — Most of the morphometric and Creek at right bank of Gran Rio, 13 km S.W. of Djoemoe meristic data in I-III. Village, depth 50-100 cm, width 5 m, bottom sand and are listed tables

mud, coll. H. Nijssen, 30-1-1967; — ZMA 106.334, seven, The number of thoracic scutes in 5 specimens 3 $ S , SL 237.2-272.5 mm (one deposited in BMNH from the Suriname River basin is 6 on both sides 1982.4.7:7), 4 of undetermined sex, SL 206-265.3 mm, district creek bank of Gran 4 km in both sides in and in two Brokopondo, at right Rio, one, 7 on two, 8/7

N.E. of N.E. part of Awadam ( = Awaradam) Fall, depth specimens. One specimen from the Saramacca width 6 bottom 30-150 cm, m, running water, sand, coll. River basin has 8/7 thoracic scutes. Two speci- H. Nijssen, 31-1-1967; — ZMA 106.339, four, SL 112.7- from the Nickerie River basin 6 227.9 mm, district Brokopondo, Awara Creek at right bank mens have

of Suriname River, 1.5 km S. of Botopasi Village, depth thoracic scutes on both sides, like the holotype 80-150 cm, width 8 m, running water, bottom sand, coll. from the River (Corantijn River basin). — Sipaliwini H. Nijssen, 18-111-1967; ZMA 106.336, five, 4 $$,

SL 1 of undetermined with about five broad 243.8-263.7 mm, sex, SL 243.8 mm, Specimens transverse

district = Creek left Brokopondo, Parwapa ( Paba) at bank bands the dorsal fin in posterior to last ray occur of Suriname River, 2.5 km N. of Botopasi Village at Foe- the rivers Suriname Nickerie, toenakaba width (fig. 4), Saramacca, (= Voetoekaba) Village, depth 50-200 cm,

8 and These bands m, running water, bottom sand, coll. H. Nijssen, 20-IH- Corantijn. are not always promi-

1967; — ZMA 106.343, 245 district S, SL mm, Broko- darkest the sides and in nent, usually at many pondo, Suriname River, rapid 7.5 km N. of Botopasi Village, the specimens bands are very faint on the mid- coll. — H. Nijssen, 27-111-1967; ZMA 106.346, three, SL dorsum of the caudal in colour 76.2-139.9 mm, district Brokopondo, creek at right bank of peduncle. Variation

Kleine Saramacca River, 11 km E.S.E. of confluence with is pattern noted below, according to the different Saramacca River, depth 30-100 cm, width 4 m, running river basins. sand and coll. water, bottom stones, H. Nijssen, 27-11-1967;

— ZMA 106.342, SL 236.5 mm, district Brokopondo, Kleine

Saramacca km River, rapids 14 E.S.E. of confluence with Suriname River basin. — In ZMA 106.338, thir-

Saramacca River, 100-200 width 80 bottom in have depth cm, m, teen specimens (121.5-169.5 mm SL) usually vague, sand, coll. H. Nijssen, 28-11-1967. broad transverse bands; nine of these specimens have numer-

ZMA 106.335, SL 117.2-162.5 mm, district indistinct brownish dots the dorsum of four, Nickerie, ous small, on the

Stondansie Fall in Nickerie River, width 80 bottom sand m, head and snout. Two specimens (102.5-119 mm) are quite

and rocks, coll. H. Nijssen, — ZMA Four 5-IV-1967; 106.340, pale. specimens (75.3-86 mm) have a well-developed

seven, SL 99.1-160 mm, district creek at colour with variable dorsum of Nickerie, right pattern, brown spots on the bank of Nickerie 12 km W.S.W. of head and River, Stondansie Fall, the body, snout; prominent spots behind the base

depth 50-100 cm, width 7 m, running bottom sand, of the last dorsal fin water, ray at the intervals where in larger

coll. H. Nijssen, 5-IV-1967; — ZMA 2 106.337, seven, $ $, specimens the bands are situated. One specimen with imper-

SL 256-266.2 5 of undetermined mm, sex, SL 100.9- fect pigmentation. district 201.5 mm, Nickerie, in Fallawatra 5 km In ZMA 106.344 broad bands rapid River, transverse are present; S.W. of Stondansie Fall, width 60 bottom sand and the smaller has the m, specimen indistinct brown spots on coll. H. rocks, Nijssen, 6-IV-I967. dorsum of the snout. - 167 DE 52 1982 BIJDRAGEN TOT DIERKUNDE, (2)

colour of the well as indistinct brownish spots; the pattern of smallest specimen is reminiscent of the juveniles Meta- four in ZMA loricaria p. paucidens and of the juveniles

106.338 (Suriname River basin). broad bands; The specimen in ZMA 106.342 has transverse of the head and snout. faint brown spots on the dorsum

Nickerie River basin. — In ZMA 106.337 the speci-

all with less broad trans- mens are well-pigmented, more or

from the dorsum of verse bands and with dark brown spots

the tip of the snout, in some specimens extending posteriorly

to near the caudal fin base. The largest specimen is very

dark dark, obscuring the visibility of spots, although trans-

well Six with minute verse bands are visible. specimens a

dark brown spot just beyond the supraoccipital process.

ZMA 106.335: bands and spots present; a minute dark

brown the spot just posterior to supraoccipital process.

for have In ZMA 106.340 all except the largest specimens brown the a dark spot just beyond supraoccipital process; Fig. 5. Map of the Guianas, showing the occurrence of two all have broad transverse bands; spots are present in some subspecies of Metaloricaria paucidens Isbrücker, 1975: M. p. specimens, although all but one of the specimens are poorly paucidens in the rivers Oyapock and Marowijne (= Maroni) pigmented. (black triangles), and M. p. nijsseni (Boeseman, 1976) in the rivers Suriname, Saramacca, Nickerie, and Corantijn Sipaliwini River. — The holotype and the topo- T indicates the (black spots). type-locality. five broad bands typical paratype (RMNH 27496) have vague

and the head. on the caudal peduncle vague spots on In ZMA 106.339 the largest specimen has posteriorly

broad bands and it has distinct on the dorsum brown — spots Corantijn River basin. The five specimens in

of the anterior to the base of the last dorsal fin ray, body RMNH 27497 are rather pale (the larger specimens are dorsum of the head and snout. One speci- extending on the recorded above, in comparison with specimens from the

men has the broad bands, no One speci- (277.7 mm) spots. Suriname River basin). The smaller ones (115 and 127 mm)

is almost without for men (129.2 mm) pigment, except have five vague broad transverse bands.

brown pigment on the caudal fin. One specimen (112.7 mm)

and bands. has distinct brown spots rather narrow transverse

All specimens in ZMA 106.336 have broad transverse

bands. The male, 243.8 mm, with minute faint brown spots REFERENCES

base of dorsal fin anterior to the the last ray, extending on

the dorsum of the head and snout; a specimen with the BAILEY, R. M. & J. N. BASKIN, 1976. Scoloplax dicra, a

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the base of the last dorsal fin ray, extending anteriorly to Pap. Mus. Zool. Univ. Michigan, 674: 1-14.

the the The Loricariinae of scutes posterior to supraoccipital process. BOESEMAN, M., 1971. "comb-toothed"

The ZMA 106.334 all have broad reflections tendencies specimens in transverse Surinam, with on the phylogenetic bands. within the family Loricariidae (Siluriformes, Siluroidei).

The specimens in RMNH 27501 are pale, the smaller one Zool. Verh. Leiden, 116: 1-56, pis. 1-8.

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mens in RMNH 27500 (231-248 mm; Saramacca River 1794), with designation of the lectotype and restriction basin) and in the larger specimens in RMNH 27497 (180- of its type locality (Pisces, Siluriformes, Loricariidae).

203 mm; Corantijn River basin). Bijdr. Dierk., 41 (1): 10-18.

The specimens in RMNH 27499 all have transverse 1972. The of the South American catfish vague , identity broad bands the caudal and on peduncle spots on the head. Loricaria cataphracta Linnaeus, 1758, with redescrip-

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specimens are rather pale; transverse bands are present, as cariidae). Beaufortia, 19 (255): 163-191. 168 I. J. H. ISBRUCKER & H. NIJSSEN - METALORICARIA PAUCIDENS

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de la and the néotropicaux, avec un catalogue critique sous- Pterosturisoma n. gen., including description

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Received: 19 July 1982