The South American Plated

Aspidoras R. von Ihering, 1907,

with descriptions of nine new species from Brazil

(Pisces, Siluriformes, )

by

H. Nijssen & I.J.H. Isbrücker

InstituteofTaxonomic Zoology, University ofAmsterdam, The Netherlands

Abstract as also Callichthys and Hoplosternum, by having

two pairs of nuchal plates between the occipital

catfish The South American callichthyid genus Aspidoras R. and the base of the dorsal; the occipital plate

von Ihering, 1907, is redefined on the basis of thirteen nearly truly hexagonal, the posterior angle not nominal species. Morphological differences with the related much produced. The head is rather elevated, not genus Lacépède, 1803, are discussed. Two species depressed, rounded in front and somewhat com- originally described in Corydoras, viz., Corydoras raimundi and The coracoid Corydoras pauciradiatus are herein transferred to Aspidoras. pressed laterally. plates entirely

Aspidoras now consists of the following previously describ- hidden by the skin. Barbels short, not reaching

ed nominal species: (1) Aspidoras rochai R. von Ihering, 1907 the gill-openings. Type: Aspidoras rochai n. sp." (type-species of the genus), known from Brazil, Est. Ceará, Gosline (1940: 10) published the following Fortaleza, (2) Aspidoras raimundi (Steindachner, 1907) from diagnosis of Aspidoras: "Head compressed. AspidorasBrazil, Est. Maranhão, Rio Parnaíba system, (3) form of lakoi P. de Miranda Ribeiro, 1949, from Brazil, Est. Minas Lower lip reverted to a single pair

Gerais, Rio Paranaíba system, and (4) Aspidoras pauciradiatus barbels beside the rictal barbels [his fig. 3], Rictal (Weitzman & Nijssen, 1970) from Brazil, Est. Goiás, Rio barbels reaching about to level of gill opening. Araguaia system. Eye more or less superior. Suborbital naked. Nine new species from Brazil are described herein: (1) Fontanel small, roundish, its length equal to Aspidoras albater from Est. Goiás, Rio Tocantinzinha, (2)

Aspidoras brunneus from Est. , Serra do Ron- about half the diameterof the eye. Supraoccipital

cador, (3) Aspidoras carvalhoi from Est. Ceará, Guaramiranga, forming a broad, short triangle posteriorly.

from Est. Ver- (4) Aspidoras eurycephalus Goiás, Córego Nuchal plates barely meeting along the midline melho, (5) from Est. Mato Grosso, between the and the supraoccipital azygous pre- Córrego Corguinho, (6) Aspidoras maculosus from Est. Bahia, dorsal Abdomen between fins from Est. plate. pectoral Rio Itapicurú system, (7) Aspidoras menezesi Ceará, covered with flesh. Dorsal its Rio Salgado system, (8) Aspidoras poecilus from Est. Mato completely 1,7,

Rio and from Est. Rio base Grosso, upper Xingu, Goiás, Araguaia, somewhat shorter than its distance from

and (9) Aspidoras spilotus from Est. Ceará, a tributary of Rio adipose. Caudal forked. One species; north- Acaráu, and from Cachoeira do Gusmão. In addition to these eastern Brazil." In his key to the genera of the nominal species some material is described and figured under family Gosline cit.: distin- of aff. These Callichthyidae, (op. 5) the provisional designation Aspidoras sp. poecilus.

another but for- guishes Aspidoras from both Corydoras and specimens may represent species, are not mally named. Brochis Cope, 1872, by the following characters:

"Foremost plates of upper lateral series, i.e.,

nuchal plates, meeting along the midlinebetween

INTRODUCTION occipital and dorsal; abdomen between pectoral

bases entirely covered with flesh; fontanel small,

R. von Ihering published a bilingual original roundish." What Gosline called Aspidoras rochai

of the of which diagnosis Aspidoras, English part (the same material identified as A. rochai by Ellis, reads is (1907: 30): "This new genus apparently 1913) actually represents a new species, here

related but from closely to Corydoras, differing it, described as Aspidoras maculosus.

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Knaack (1966: 365), while describing Corydoras [IU1 Indiana University (material now in CAS).

MNRJ Museu Nacional, Universidade Federal do Rio pygmaeus as a new species, noted that: "Nach der de Janeiro, Rio de Janeiro. Z. noch im z. allgemeinen benutzten Systematik MZUSP Museu de Zoologia da Universidade de Sao der Callichthyiden von Gosline (1940) miiRte Paulo, Sao Paulo. diese Art der Gattung Aspidoras zugeordnet NMW Naturhistorisches Museum, Vienna.

werden. Auf Grund vorliegender Untersuchungs- RMNH Rijksmuseum van Natuurlijke Historie, Leiden.

[SU1 Stanford now ergebnisse lafet sich diese Gattung nicht aufrecht University (material in CAS). UMMZ University of Michigan, Museum of daK die Zoology, erhalten, so Art pygmaeus dem Genus Ann Arbor, Mich. Corydoras zugeschrieben wurde. Eine ausfuhr- USNM National Museum of Natural History, Washing-

liche Diskussion und iiber den Ein- D.C. Begriindung ton,

der wird in einer ZMA Instituut voor zug Gattung Aspidoras spateren Taxonomische Zoologie (Zoolo- gisch Museum), Amsterdam. Arbeit erfolgen." To our knowledge, Knaack bd body depth, measured from the anterior edgeof never published this intended paper. the azygous predorsal scute vertically to the

Hoedeman (1952) — based on Gosline, 1940 — ventral edge of the ventrolateral body scute(s) united Brochis, and in the Aspidoras, Corydoras (where in Corydoras the coracoid is exposed); in

adult females the subfamily Corydoradinae Hoedeman, 1952 bearing ripe eggs bulging belly

is not included;expressed as aratio of si. (forming together with the Callichthyinae Gill, width, bw body measured between point just an- 1872, the family Callichthyidae Gill, 1872). He terior to the pectoral spine insertions; expressed split the Corydoradinae into two tribes, the as a ratio of si.

with Corydoras and Brochis, and the width of "coracoid Corydoradidi ca area", measured between

Aspidoradidi with Aspidoras. We prefer to omit the ventral edges of the anteriormost ventrolat-

discussion of the higher classification (above eral scutes; expressed as a ratio of hi. dbs number of dorsolateral body scutes (except the generic level) within the family Callichthyidae small scutes at the posterior end of the until suitable comparative anatomical investiga- caudal peduncle).

tions have been least of the caudal accomplished. dcp depth peduncle; expressed as

ratio of hi. The majority of the species we recognize a

within h! head length, measured from the snout Aspidoras show greater similarities than tip (ethmoid) to the highest point of the gill differences. Four hundred specimens were avail- opening; expressed as a ratio of si. able for examination. We ascribed them to lbo length of the bony orbit, horizontally measured; thirteen nominal We realize that the species. expressed as a ratio of hi.

results presented in this publication have not to Ids length ofthe dorsal fin spine, measured with the

be considered clear-cut solutions of the problems spine in vertical position from the junction of the bases of the dorsal spine and the predorsal we were faced with. Many of the specimens at scute to the spine tip; expressed as a ratio of si. hand were collected a long time ago, and we have Ips length of the pectoral spine, measured with the to await freshly collected material to describe the spine pressed along the body from the articula- minor details of the colour The tion point of the its patterns. mor- spine to tip; expressed as a

phometric and meristic data represented in the ratio of si. pas number of pre-adipose scutes, is the number of bar diagrams (figs. 17 through 19) were taken middorsal scutes anterior to the adipose fin from a limited number of adult specimens, for spine, which is in fact a modified middorsal are hence not useful many specimens juvenile, scute thoughnot included in the count. for comparison with adult specimens of other si standard length, measured from the snout tip species. (ethmoid)to the junction of the posterior edges

of the last counted scutes of the caudal pe-

duncle; expressed in mm tothe nearest tenth, ABBREVIATIONS USED AND DEFINITIONS OF TERMS sn snout length, measured from the snout tip CAS California Academyof Sciences, San Francisco, (ethmoid) to the nearest anterior edge of the

Calif. bony orbit; expressed as a ratio of hi. [CM) vbs Carnegie Museum (material now in FMNH). number of ventrolateral body scutes (except the DNOCS Departamento Nacional de Obras Contra as small scutes at the posterior end of the caudal

Secas, Fortaleza, Ceara. peduncle).

FMNH Field Museum of Natural History, Chicago, 111. wi least width of the interorbital, is the least IRScNB Institut Royal des Sciences Naturelles de Bel- distance between the dorsal borders of the orbit

Brussels. of each gique, side; expressed as a ratio ofhi.

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ACKNOWLEDGEMENTS more dorsal fin rays (10-17 in Brochis against 6-8

(usually 7) in Corydoras). of like For the loan and/or exchange specimens we to express Aspidoras comprises small sized species, up to our gratitude to the following colleagues: Dr. R. M. Bailey & 41.8 mm standard length (Aspidoras menezesi n. Dr. R. R. Miller (UMMZ); Dr. M. Boeseman (RMNH); Dr.

Dr. R. A. Braga (DNOCS); Dr. H. Britski (MZUSP); A. sp.), with small eyes and short dorsal and pecto-

Leitao de Carvalho (MNRJ); Dr. W. N. Eschmeyer & Miss P. ral spines. In habitus, all but one species (Aspi- Sonoda Dr. J. Dr. J. (CAS); Gery, Saint-Cyprien, Dordogne; unlike the doras poecilus n. sp.) are majority of P. Gosse & Mr. E. Walschaerts (IRScNB); Dr. R. K. Johnson Corydoras and Brochis species roundish rather (FMNH); Dr. P. Kahsbauer (NMW); and Dr. S. H. Weitzman than in their transverse the triangularly depressed (USNM). Mr. L. A. van der Laan (ZMA) made photo-

graphic illustrations. Figs. 2 and 3 were drawn by Mrs. M. P. head shape.

Bakry (USNM); fig. 5 by Dr. S. H. Weitzman. Dr. Weitzman Too much emphasis has hitherto been laid also read the and discussed 6 and 25 kindly manuscript (in lit., of the nuchal upon the characteristic plates the of the An November, 1975) status genus Aspidoras. meeting along the midline between occipital agreement with our opinion that Aspidoras is distinct from process and dorsal fin spine, to distinguish Aspi- Corydoras is not implied. doras from Corydoras. The position of the nuchal

insuf- plates (i.e., meeting or not meeting) is an

Aspidoras R. von Ihering, 1907 ficient character to distinguish Aspidoras from

in of the Corydoras, as some species Aspidoras Aspidoras R. von Ihering, 1907: 30 and 31 (original nuchal plates are separated along the midline by diagnosis, in English and Portuguese; type-species, by the like in most original designation and monotypy, Aspidoras rochai R. von supraoccipital process (fig. 16n),

Ihering, 1907). — not all — species of Corydoras.

Little is known of the evolution of the Cal-

Aspidoras is easily distinguished from all other lichthyidae and therefore it is speculative to

callichthyid genera (Corydoras, Brochis, Dianema, discuss phylogenetic relationships. For example,

Callichthys, Hoplosternum) the of of by possession we do not know whether the presence a two cranial fontanels. The anterior (frontal) fontanel in supraoccipital (pit) Aspidoras repre- fontanel round or oval in slightly shape sents a retained primitive character shared with

the roof of the whereas always penetrates skull, unknown ancestral callichthyids or that it is a the posterior fontanel is closed in (supraoccipital) specialized character in a process of closing the small roundish adult specimens, leaving a shallow is skull. If the character primitive (the presence

is considered pit. of a frontal and parietal fontanel In the 400 examined of thirteen specimens primitive for ostariophysan fishes), then it is the species we assign to genus Aspidoras a hardly defendable to recognize Aspidoras as a similar of basically structure the skull roof was taxon different from Corydoras according to the The of this character is found. variability small. Hennigian cladistic approach. On the other hand, None of the other Callichthyidae show this none of the known callichthyid generashares, this and intermediate structure no structure was character, so it might be considered a specialized

found in the 100-odd of the related species genus character. We are unable at the moment to use

Corydoras. this character for making phylogenetic inferen-

Corydoras, Brochis, and Dianema a useful assemble possess ces, but certainly it is to the single, fontanel much larger and far thirteen which the open species possess supraoccipital more elongate than the frontal fontanel in Aspi- pit in a separate genus. doras — with a commissural bar anteriorly.

Callichthys and have a roundish Hoplosternum Etymology. — frontal with fontanel a foramen anterior to the Not explained by R. von Ihering. Most probably commissural bar and a small foramen posterior to Aspidoras is derived from the Latin "asper", it. and from meaning rough, harsh, uneven, the

is most related Greek is Aspidoras closely to Corydoras 8opa, meaning skin, hide. There a well- 1803 Brochis Lacepede, and Cope, 1872. Brochis known generic name in Siluriformes, Doras La - differs from Corydoras among others by having cepede,1803.

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Table I. Morphometric characters in mm to the nearest tenth, and counts ofthe primary type-specimens of the species of Aspi-

doras.

si Ids bd bw hi sn lbo wi ca D A P C dbs vbs lps dcp P, 2 pas

rochai 39.9 10.6 10.4 — — 10.2 5.4 1.9 4.5 4.5 5.5 1,7,i i,5 i,6 1,9 7/7 27 24 6

raimundi 25.5 6.7 6.0 4.0 4.5 6.4 3.1 1.8 3.0 2.0 3.6 I,7,i i,6 ii,5 1,9 7/7 25 23 4

lakoi 33.4 10.4 9.1 3.2 5.1 9.6 4.8 1.2 4.6 3.2 4.9 1,7,i i,6 i,5 1,9 in 26 24 11

pauciradiatus 23.2 6.8 5.9 4.5 5.5 6.2 2.5 1.7 2.8 3.5 2.9 1,6,i i,5 i,6 1,7 7/7 23 20 2

albater 35.6 9.0 8.9 4.4 6.5 9.9 5.5 2.0 4.0 3.9 4.8 1,7,i i,6 ii,5 1,8 7/7 26 23 5

brunneus 20.9 5.4 5.3 3.0 3.6 5.8 3.1 1.5 2.4 2.2 3.0 1,7,i i,6 ii,5 1,9 7/7 26 23 4

carvalhoi 25.4 7.4 6.8 3.9 4.9 6.6 3.4 1.3 3.2 3.1 3.8 I,7,i i,5 ii,5 1,9 IP 25 22 4

eurycephalus 29.5 8.2 7.5 2.9 3.3 8.0 4.3 1.5 4.2 3.0 4.5 1,7,i i,5 ii,5 1,9 7/7 26 23 4 fuscoguttatus 32.0 9.2 8.5 4.7 4.9 8.6 4.5 1.4 4.0 4.0 4.7 1,7,i i,5 i,5 1,8 7/7 27 24 3

maculosus 37.2 9.7 8.7 4.8 5.5 9.5 5.1 2.3 3.8 5.8 4.8 I,7,i i,5 ii,5 1,8 7/7 26 23 3

menezesi 41.8 11.2 10.3 5.8 7.5 10.9 5.9 2.4 4.1 4.7 5.5 I,7,i i,5 ii,5 1,9 7/7 26 23 6 poecilus 29.2 8.3 7.1 5.4 5.8 7.8 4.1 1.9 3.7 2.4 4.2 I,7,i i,5 ii,5 1,8 7/7 25 22 6

spilolus 34.0 8.2 8.1 4.2 5.5 9.3 5.1 1.9 3.2 3.5 4.1 I,7,i i,5 i,5 1,9 7/7 25 22 3

A = anal fin; C = caudal fin; D = dorsal fin; P, = fin; P = fin. pelvic 2 pectoral

KEY TO THE SPECIES OF ASPIDORAS b. Body with irregular oblique brown marbling; length

dorsal 6.3 8.9 in si aff. spine to . Aspidoras sp. poecilus

la. Snout 2.5 in less with more less roundish dorsal length or more hi; length pectoral spine c. Body or spots; length

than 4.5 in si: number of dorso- and ventrolateral body spine more than 5.8 in si 10

dorsal fin with and 6 soft last 10a. Dorsal fin with a dark scutes 23/20; a spine rays, usually conspicuous, triangular

ray split to its base blotch . . . Aspidoras raimundi (Steindachner, 1907)

Aspidoraspauciradiatus (Weitzman& Nijssen, 1970) b. Dorsal fin without a conspicuous, triangular dark

b. Snout 2.1 less blotch length or in hi; length pectoral spine more 11 than 4.5 in si; number of 11a. Length dorso- and ventrolateral body pectoral spine more than 8.0 in si; interorbital

scutes 25/22 or more; dorsal fin with a spine and 7 soft width less than 2.0 in hi; length dorsal spine more than

last to its base 2 9.5 in si rays, ray split Aspidoras eurycephalus i- sp.

2a. Dorsolateral body scutes with a longitudinal solid dark b. Length pectoral spine less than 8.0 in si; interorbital

brown band brunneus width than 2.0 in dorsal Aspidoras n. sp. more hi; length spine less than

b. Dorsolateral body scutes without longitudinalsolid dark 9.5 in si 12

brown band 3 12a. Length dorsal spine more than 7.3 in si; length pectoral

3a. Body with about 4 distinct oblique bars spine more than 5.7 in si; body depth 3.8 to 4.0 in si; body

albater width 4.2 4.4 in 3 4 - Aspidoras n. sp. to si; to pre-adipose scutes b. Body with other colour without colour maculosus markings or Aspidoras n. sp.

markings 4 b. Length dorsal spine less than 7.3 in si; length pectoral

4a. Body with distinct dark midlateral blotches 5 spine less than 5.7 in si; body depth 3.3 to 3.8 in ... si; body

b. Body without distinct dark midlateral blotches 6 width 3.8 to 4.2 in si; 5 to 6 pre-adipose scutes

5a. Lengthbony orbit 5.3 or more in hi; width 3.8 in menezesi n. body si; Aspidoras sp. 6 pre-adipose scutes

Aspidoras rochai R. von Ihering, 1907 Aspidoras rochai R. von Ihering, 1907 b. Length bony orbit 5.0 or less in hi; body width 3.9 or (figs. 1, 15a-b, 16a, 17-20) more in si; 3 to 4 pre-adipose scutes

Aspidoras spilotus n. sp. Aspidoras rochai R. von Ihering, 1907: 30-34 (original 6a. without colour carvalhoi Body pattern Aspidoras n. sp. description; type-locality: "Fortaleza, capital of State of b. with colour Body a pattern 7 Ceard"; lectotype in Departamento de Zoologia, Secre- 7a. orbit 6.5 in Length bony or more hi; 6 or (usually) more taria da Agricultura, Sao Paulo, MZUSP 2195); Eigen- pre-adipose scutes mann, 1910: 402 (listed); A. de Miranda Ribeiro, 1911: 155- Aspidoras lakoi P. de Miranda Ribeiro, 1949 156 (description, after R. von Ihering); A. de Miranda b. Combination of orbit and number length bony of pre- 1918: 720 Ribeiro, (listed; apparently the syntypes; Forta- adipose scutes not as in 7a 8 leza); Fowler, 1954: 46 (references, in part); Britski, 1969: 8a. marbled with brown Body and with some rows of small 206 (description; designation of the lectotype). roundish dark brown spots (lacking in juveniles) . . .

Material examined. — One - Aspidorasfuscoguttatus n. sp. (lectotype, designatedby Britski,

b. Body marbled with brown, without rows of small 1969: 206), MZUSP 2195, si 39.9 mm, Brazil, Est. Ceara,

roundish dark brown spots 9 Fortaleza, 03°45' S, 38°35' W, coll. F. Dias da Rocha, 1-1905; with 9a. Body one irregular oblique brown marbling; length (paralectotype), MZUSP 5300, si 37.8 mm, same data as dorsal spine 5.2 to 5.4 in si Aspidoras n. poecilus sp. lectotype.

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Fig. 1 Aspidoras rochai R. von Ihering, 1907. Lectotype.

Description. — and somewhat irregular band from the nake to Morphometric and meristic data in table I and the caudal, being united in front with that of the

figs. 17 through 19. Both the lecto- and para- other side. Pectoral and ventral fins plain; anal

in condition. The dorsal fin with several which form dorsal lectotype are a poor spots a cross-line;

spine is broken in the lectotype. Dorsal fin rays dark at the base, light in the middle and as it are broken off near the base in both specimens, seems, with dark subterminal markings; caudal whereas the caudal fin is almost completely lost with 4-6 (?) oblique series of little spots." in both Also the and anal In footnote 32 specimens. pectoral a on page R. von Ihering states: fins in the here the full of the are badly damaged, although paralec- "I cannot give description

observe that and totype we the middle pectoral fin dorsal caudal fins as they are greatly damag- rays reach over the pelvic fin base to about the ed in both typical specimens." tip of the first (unbranched) pelvic fin ray. Only

the pelvic fins are still in good shape in the Etymology. — lectotype. In honour of Mr. Francisco Dias da Rocha (R.

Snout in front of nostrils naked. Two short von Ihering, 1907: 34 and 35). mental barbels, two pairs of rictal barbels, longer

(upper) rictal barbel reaching to or a little Aspidoras raimundi (Steindachner, 1907) vertical from of beyond a anterior margin eye. (figs. 2-3, 15c, 16b, 17-20) Inner edge of pectoral fin spine moderately serrated (figs. 15a-b). Anterior fontanel short- Corydoras raimundi Steindachner, 1907: 84-85 (original oval (fig. 16a). Skin of intercoracoid area with description; type-locality: "... in dem Bachchen, welches isolated minute de- numerous ossifications, bei Victoria in den Rio Parnahyba mundet."; lectotype in creasing in number and size posteriorly. Naturhistorisches Museum, Vienna, NMW 61110); Eigen-

mann, 1910: 403 (listed); Regan, 1912: 215 (description,

Colour in alcohol — for some 1). Except on (fig. after Steindachner; in key p. 210); Ellis, 1913: 409

in after very faint, indefinite brownish shading along the (listed; key, Steindachner, on p. 400); Gosline,

has 1940: 21 (listed; in key on p. 14); Gosline, 1945: 75 (listed); midline of the body, the colour pattern Fowler, 1954: 66 (listed); Nijssen, 1970: 58 (listed; placed completely disappeared from the lectotype and within Corydoras “barbatus-group”). the paralectotype. R. von Ihering (1907: 32 and Corydoras raymundi; Van der Stigchel, 1946 & 1947: 130- described the colour in the of the 34) English part 131 (misspelling of name; description of "type", which is

listed 5 original description as follows: "General colour on p. as "holotype"). of body dark-brown above, light below; several Material examined. — One (lectotype, by present designa- rounded and light spots on operculum flanks; tion), NMW 61110 (ex NMW 46794), si 25.5 mm, Brazil, Est.

brook Rio Parnaiba above the lateral line runs, parallel to it, a light Maranhao, at mouth of emptying into at

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Alto Parnaiba (formerly Victoria), 09°08' S, 45°56' W, coll. F. fin spine. Inner edge of pectoral fin spine strong- Steindachner, 26/29-VI-1903; sixty-two (paralectotypes), ly serrated (fig. 15c). Anterior fontanel elongate NMW 46792 (one), NMW 46794 and 46795 (fifty-one), ZMA oval (fig. 16b) in the lectotype and most of the 110.480 (nine, ex NMW 46794), RMNH 7962 (one, this other short-oval in some para- specimen has erroneously been considered "holotype" by paralectotypes, Skin of intercoracoid with Van der Stigchel, 1946 & 1947: 5), si 12.7 to 32.6 mm, same lectotypes. area data as lectotype. several isolated minute ossifications, decreasing

in number and size posteriorly.

Description. — Colour in alcohol (figs. 2-3). — Steindachner

Morphometric and meristic data in table I and (1907: 85) describes the colour pattern as: "3

figs. 17 through 19. Dorsal and pectoral spines Reihen langlicher Fleckchen von grauvioletter than Hohendrittel distinctly shorter adjacent rays. Snout in Farbung langs der beiden oberen

Schwanzflosse front of nostrils naked. Two short mental barbels, des Rumpfes. quer gebandert.

two of rictal which reach Eine fast scharf Binde pairs long barbels, may schwarzliche, abgegrenzte

(as in the lectotype) well beyond base of pectoral langs der Hohenmitte der Ruckenflosse, zu-

Fig. 2. Aspidoras raimundi (Steindachner, 1907). Lectotype, male.

Fig. 3. Aspidoras raimundi (Steindachner, 1907). Paralectotype, female,NMW 46794, sl 32.6 mm.

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re-examined at hand. there weilen nicht ganz bis zum hinteren Flossenrande we Anyhow, was no reichend." holotype selected for Corydoras raimundi, as Van

Ground colour of body and head pale tan. der Stigchel assumed. Probably, Van der Stigchel

that the from Small, irregular brown spots distributed over thought species was described a body, operculum and cleithrum. In some speci- single specimen. mens spots tend to be arranged into a longitu- but dinalpattern. No mask present melanophores

dense between eyes and over supraoccipital Aspidoras lakoi P. de Miranda Ribeiro, 1949

dark brown bone, producing a colour on poste- (figs. 4, 15d, 16c, 17-20) rior dorsum of head. Dorsum of snout pale tan to

Aspidoras lakoi P. de Miranda Ribeiro, 1949: 143-145, 1 pale brown. Belly pale tan, without dark pigment. fig. (original description; type-locality: "Pequeno c6rrego na Barbels all pale tan to brown. floresta do Grotao, Fazenda da Cachoeira, Municipio de Dorsal fin unpigmented along base and tip, Passos, Estado de Minas Gerais"; holotype in Museu less with a prominent dark brown, more or Nacional, Rio de Janeiro, MNRJ 5292); P. de Miranda triangular blotch, originating at about halfway Ribeiro, 1959: 8 (listed; holotype and topotypes: Est. the dorsal spine. Adipose fin spine usually dark Minas Gerais, Corrego na floresta do Grotao, fazenda da Cachoeira, Passos). brown, and some dark brown pigment about this

Caudal fin with Material examined. — One (holotype), MNRJ 5292, si 33.4 spine on the membrane. rays

mm, Brazil, Est. Minas Gerais, Rio Parana system, a tributary minute dark brown spots, forming narrow, ir- of Rio floresta do Grande, pequeno corrego na Grotao, regular, more or less vertical to oblique bars. Fazenda da Cachoeira, Municipio de Passos, coll. C. Lako, and anal fins without Pectoral, pelvic, pigment. IX-1945; twenty-three (topotypes), MNRJ 5293 (twenty-one),

ZMA 113.565 (two), si 14.9 to 39.7 mm, coll. C. Lako, 1945.

Etymology. —

Unknown, perhaps named after a Brazilian as- Description. — sistant (Dr. P. Kahsbauer, in lit.). Morphometric and meristic data in table I and

figs. 17 through 19. Dorsal and pectoral fin spines

Nomenclaturalnote. — distinctly shorter than adjacent rays. Snout in

Steindachner (loc. cit.) did not state the exact front of nostrils naked. Two short mental barbels, number of specimens he had at hand while two pairs of long rictal barbels, reaching the gill originally describing this species. We are quite openings ventrally. Inner edge of pectoral spine

sure he had at least the sixty-three syntypes that faintly serrated (fig. 15d). Anterior fontanel elon-

Fig. 4. Aspidoras lakoi P. de Miranda Ribeiro, 1949. Holotype.

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oval in the and de gate some specimens, slightly rhomboidal collected holotype topotypes (P.

in others (fig. 16c). Skin of intercoracoid area Miranda Ribeiro, 1949: 143).

naked without ossifications.

in — P. de Miranda Colour alcohol (fig. 4). Aspidoras pauciradiatus (Weitzman & Nijssen, the Ribeiro (1949: 144) describes colour pattern 1970) as: "O colorido fundamental e isabelino, sendo (figs. 5, 15e, 16d, 17-20)

que as placas, especialmente nas extremidades,

F-223.00 mostram maculas castanhas que, vistas em con- “Corydoras U-6" Axelrod et al., 1967: (colour

photograph). junto, parecem formar uma tarja ao longo da Corydoras pauciradiatus Weitzman & Nijssen, 1970: 129- linha lateral e maculas irregularmente dispostas 131, figs. 5, 6e (original description; type-locality: "Brazil, manchada sobre o corpo; cabecja irregularmente Rio Araguaia near Aruana (14°58' S, 51°04' W), Est. de castanho, mais carregado o focinho. para Goias"; holotype in United States National Museum, sendo Dorsal e adiposa escuras na base, que a Washington D.C., USNM 191625). primeira com pintas castanhas sobre os raios; Material examined. — One (holotype), USNM 191625, si 23.2 anal com uma macula castanha na base, apos o mm, Brazil, Est. Goias, Rio Araguaia near Aruana, 14°58' S, tendo demais manchados primeiro raio, os em 51°04' W, coll. H. R. Axelrod, 1960; one (paratype), USNM

204363, si 22.6 mm, same data as ZMA sua metade, formando uma tarja naquela regiao. holotype; two, 113.586, si 20.1-21.4 mm, aquarium specimens, donated by F. Os raios de caudal, com manchas castanhas, Wolter, X-1970; one, ZMA 113.587, si 29.4 mm, aquarium arrumadas paralelamente em seis filas." specimen, donated by R. Suhr, specimen was imported in Ground colour Holotype. yellowish. Large, 1963, died XII-1968. irregular, alternating (originally dark?) brown blotches all over the sides of the body. Adipose Description. — and of membrane dark brown. Dorsal spine part Morphometric and meristic data in table I and fin with small brown dots. Dorsum of some figs. 17 through 19. Two pairs of rictal barbels fin brownish. Pelvic fins Anal fin pectoral plain. and one pair of mental barbels. Inner edge of

basal followed tan with rather large dot, by a pectoral spine strongly serrated (fig. 15e). Fon-

which is followed a dark tan zone, by oblique tanel round, 0.7 mm in diameter in holotype and Caudal fin zone. Tips of anal fin rays pigmented. Skin of paratype, enclosed by frontals (fig. 16d). broken off for the according to greater part, but, intercoracoid area naked. the rather good photograph of this specimen

Colour in alcohol (fig. 5). — Ground colour of published in the original description, with many body and head tan. No mask but small brown dots, arranged into about four more pale present dense between and less lines. melanophores eyes over or regular vertical brown bone, producing a dark Topotypes. Ground colour of body and head supraoccipital dorsum of head. Dorsum of colour on posterior pale tan. Caudal fins greatly damaged in all with to 8 moderate sized brownish. snout pale brown 7 spots specimens. Bony parts of head pale with end of at of in a triangular pattern one triangle Body marbled with light brown in a sort Small brown in of snout tip. spots (as fig. 5) arranged checkerboard pattern in some the specimens. be over sides of and head. Spots tend to Dorsal fin unpigmented in some specimens, body

in a linear each brown the in arranged longitudinal pattern, with some light spots on soft rays of row occurring on 6 to 10 scutes. Posterior end other specimens, the membrane between spine toward between each row converges junction and first ray pigmented with brown in some dorso- and ventrolateral body scutes. Thus pos- specimens. Adipose fin sometimes with a faint dorsal of of the teriorly, rows spots converge ventrally small light brown dot near the base spine. toward ventral rows of which dorsum of fin spots converge A few spots may occur on pectoral The effect resembles a series of but this fin is with dorsally. vaguely rays, mostly evenly pigmented This best be posteriorly pointing cheverons. can light brown, and in some specimens it is unpig- of seen in a live colour photograph paratype on mented. Pelvic fins plain. Anal fin base with or of " U-6" in Axelrod page F-223.00 Corydoras et without a faint brown blotch. al. (1967). Belly pale tan, without dark pigment.

Etymology. — Dorsal rictal barbels brown, ventral rictal barbels

Named in honour of Mr. Carlos Lako, who white.

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Fig. 5. Aspidoraspauciradiatus (Weitzman & Nijssen, 1970). Holotype.

eleven MZUSP 12992 (four), Dorsal fin with Brasil, 1-1974; (paratypes), a large basal black blotch, on USNM 213540 (five), ZMA 113.592 (two), si 27.5 to 33.9 rays and membrane and one or two distal narrow mm, same data as holotype. horizontal bands. Pigment of these bands con-

fined fin to fin rays. Adipose with some pigment Description. — dorsally in both type-specimens; adipose spine Morphometric and meristic data in table I and with black pigment. Pectoral and pelvic fins figs. 17 through 19. Dorsal and pectoral fin spines

without dark pigment. Anal fin with one or two distinctly shorter than adjacent rays. Snout in rows of pigment, confined to the fin rays and front of nostrils naked. Two short mental barbels,

forming a band. Caudal fin with about five rows two pairs of rictal barbels, the longest (dorsal) of narrow black bars, third from anterior band reaching to or little beyond a vertical from the being darkest. Distal bar almost not distinguish- of posterior margin eye. Inner edge of pectoral able. Pigment confined to fin rays. fin spine finely serrated (fig. 150- Anterior fon- In two of the three additional specimens (ZMA tanel round (fig. 16e). Skin of intercoracoid area 113.586 and 113.587) the distal half of the pelvic with numerous isolated minute ossifications. fin rays bear some blackish pigmentation; the three specimens have dark pectoral spines. Colour in alcohol (fig. 6). — Ground colour of

body and head pale tan. Four large, irregular,

almost solid black dark brown Etymology. — oblique, or blotch-

and From the Latin "paucus" meaning few, and es on body, some smaller, more or less

isolated dots along dorsum. Dorsum of head dark "radius" meaning ray of the sun or spoke of a

brown, and dorsum of snout marbled wheel, hence rays of a fin. In reference to the few operculum dorsal with tan and dark Cleithrum and pectoral fin rays (Weitzman & Nijs- grey. even grey

for white area just superior to pectoral fin sen, 1970: 130). except spine base.

fin Base of dorsal rays with grey pigment in Aspidoras albater n. sp. most specimens; dorsal fin rays with one or two (figs. 6, 15f, 16e, 17-20) horizontal rows of dots. fin grey Adipose spine

grey, adipose fin membrane unpigmented. Cau- Material examined. — One (holotype), MZUSP 12991, si 35.6

dal fin (damaged in all specimens) with three or mm, Brazil, Est. Goids, Rio Tocantinzinha near Sao Joao da few rather broad vertical Alianpa, 14°46' S, 47°30' W, Rio Tocantins system, coll. G. a more or irregularly

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6. albater Fig. Aspidoras n. sp. Holotype.

curved bars. Pectoral and pelvic fins unpig- the road Chavantina-Casximba, coll. E. J. Fittkau, 17-VIII-

anal fin with 1965; five (paratypes), ZMA 109.380 (four), USNM 213569 mented; some faint pigment forming to (one), si 19.4 21.3 mm, same data as holotype. diffuse a spot variably extending to about halfway

the rays.

Description. —

and meristic data in table I and Etymology. — Morphometric 17 19. The The specific name is derived from the Latin figs. through six specimens used for

"albus" meaning white, and from the Latin this description are apparently all juveniles. Dor- sal and fin "ater" meaning black, in allusion to the colour pectoral spines distinctly shorter than

adjacent Snout in front of naked. pattern. rays. nostrils Two short mental barbels, two pairs of long rictal

barbels, reaching nearly to the gill openings Aspidoras brunneus n. sp. ventrally. Inner edge of pectoral fin spine strong- (figs. 7, 15g, 16f, 17-20) ly serrated (fig. 15g). Anterior fontanel long oval

to slightly rhomboidal (fig. 16f). Skin of inter- Material examined. — One (holotype), ZMA 113.588, si 20.9

mm, Brazil, Est. Mato Grosso, Serra do Roncador, km 125 of coracoid area naked.

7. brunneus Fig. Aspidoras n. sp. Holotype.

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Material examined. — One MNRJ si 25.4 Colour in alcohol (fig. 7). — Ground colour of (holotype), 5230,

mm, Brazil, Est. Ceard, Agude Canabrava, Guaramiranga, body and head pale tan. Upper half of body solid R. coll. S. de Menezes, XII-1947; one (paratype), ZMA dark brown to about the midlateral line, curving 1 13.589,si 22.3 mm, same data as holotype. downwards near caudal fin base. Ventral side of

this colour area irregularly alternated with pale

Description. — tan of ventral half of body. Ossified parts of head Morphometric and meristic data in table I and dark brown, naked part of snout with irregularly figs. 17 through 19. Dorsal and pectoral fin spines distributed dark brown chromatophores. Dorsal in distinctly shorter than adjacent rays. Snout fin with a dark blotch at base of first three rays, front of nostrils naked. Two comparatively long followed by a pale tan, unpigmented area. Distal- mental barbels (about twice the length of such ly the dorsal fin rays may bear dark pigment; in barbels in most of the other Aspidoras species), some specimens this pigment is situated lower on two pairs of rather short rictal barbels, reaching the and is followed rays by an unpigmented tip. to a vertical through the centre of the eye. Inner Adipose fin spine brown in most of the speci- edge of pectoral fin spine strongly serrated (fig. mens. Caudal fin with two or three oblique dark 15h). Anterior fontanel elongate oval (fig. 16g). brown lines. The caudal fin tips are damaged in Skin of intercoracoid area with several isolated all six specimens. A rather large, dark brown minute ossifications, decreasing in number pos- blotch in posterior half of anal fin. Pelvic fins teriorly. unpigmented. Pectoral fins usually with some

dark, irregular and faint markings. Colour in alcohol (fig. 8). — Ground colour light

yellowish umber all over. This species is almost

Etymology. — completely devoid of chromatophores; only a

The specific name is derived from the Medieval few dark brown isolated minute pigment spots on

lateral concentrated Latin "brunneus" meaning dusky, dark, tawny, in the body scutes, along or

the of the allusion to the dark brown colour pattern. near posterior edge scutes.

Etymology. — Aspidoras earvalhoi n. sp. Named in honour of Dr. Antenor Leitao de (figs. 8, 15h, 16g, 17-20) Carvalho, Chefe do Departamento de Verte-

brados of the Museu Nacional, Rio de Janeiro, Aspidoras rochai [non R. von Ihering, 1907]; P. de Miranda who kindly sent us specimens of various Ribeiro, 1959: 8 (in part; listed; Brasil, Est. Ceard, Acude many loan. Canabrava, Guaramiranga). species of Aspidoras on

carvalhoi Fig. 8. Aspidoras n. sp. Holotype.

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Fig. 9. Aspidoras eurycephalus n. sp. Holotype.

Aspidoras eurycephalus n. sp. Aspidoras fuscoguttatus n. sp. (figs. 9, 15i, 16h, 17-20) (figs. 10, 15k, 16i, 17-19,21)

Material examined. — CAS si 29.5 One (holotype), 16010, Material examined. — One (holotype), MZUSP 8573, si 32.0 Brazil, Est. Goias, Rio Tocantins mm, system, Corrego mm, Brazil, Est. Mato Grosso, Rio Parana system, Corrego

Vermelho into Rio das of Rio Almas, tributary Maranhao, Corguinho, estrada da Tres Lagoas (20°46' S, 51°43' W) —

about 15° 49°30' coll. C. six S, W, Ternetz, 26-XII-1923; Concei?ao do Taboado, coll. H. Britski, 28-XII-1968;twenty-

(paratypes), CAS 31833 (five), ZMA 113.593 (one), si 20.0 to four (paratypes), MZUSP 11737 through 11755 (nineteen),

25.4 mm, same data as ZMA 113.594 holotype. (five), si 19.0 to 38.0 mm, same data as holotype.

Description. —

Morphometric and meristic data in table I and Description. — figs. 17 through 19. Dorsal and pectoral fin spines Morphometric and meristic data in table I and

shorter than Snout in figs. 17 through 19. Dorsal and fin distinctly adjacent rays. pectoral spines front of nostrils naked. Two short mental barbels, distinctly shorter than adjacent rays. Snout in

two pairs of long rictal barbels, somewhat front of nostrils naked. Two short mental barbels,

shrivelled in all specimens. Inner edge of pectoral two pairs of long rictal barbels, reaching base of

fin spine moderately serrated (fig. 15i). Anterior pectoral fin spine in some of the specimens; in

fontanel round the rictal barbels to slightly oval (fig. 16h). Skin of other specimens are slightly

intercoracoid area naked. shorter, reaching almost to the gill openings

ventrally. Inner edge of pectoral fin spine finely Colour in alcohol (fig. 9). All seven specimens serrated (fig. 15k). Anterior fontanel elongate are badly faded, no doubt as a result of more oval (fig. 16i). Skin of intercoracoid area with than fifty years of preservation. There are, how- several isolated minute ossifications. ever, light brown markings on the body, remi-

niscent of those found in Aspidoras poecilus and Colour in alcohol (fig. 10). — Ground colour of

and head tan. Dorsum and sides of Aspidoras sp. aff. poecilus. Three specimens have body pale

remains and marbled In some of caudal fin rays, with an indica- head, opercles, snout with brown.

tion of a narrow vertical or little oblique line. most specimens there is a narrow, solid brown

oblique line down forward from the running eye.

Etymology. — Dorsum and sides of body with three kinds of

dark The specific name is derived from the Greek brown markings: (1) rather large, irregular

eiipiu meaning broad, wide, and from the Greek blotches along the midline, (2) medium-sized,

KopaXri meaning head, in allusion to its wide irregular and somewhat faint blotches just below interorbital. the dorsum of the body and just above the

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10. Fig. Aspidorasfuscoguttatus n. sp. Holotype.

ventrum of the body, and (3) series of quite 394, pi. XXVI fig. 3 (materiallisted from Agua Branca, Rio

which be Zinga, and Rio Paiaia); Gosline, 1940: 10 (two of the regular, small roundish speckles, can recorded Rio in In the specimens by Ellis, 1913; Paiaia); Gosline, arranged into rows some specimens. two 1945: 74 (listed; bacia do rio Itapicuru, Bahia, Brasil); smallest specimens (si 19.0 and 22.3 mm) these Fowler, 1954: 46 (in part; references). speckles are absent or nearly absent. Upper third of lateral body scutes, just below dorsumof body, Material examined. — One (holotype), FMNH 54810 [ex CM shaded with brown. Colour yellowish pattern 3457], si 37.2 mm, Brazil, Est. Bahia, Rio Paiaia, into

highly variable in minor details. headwaters of Rio Itapicuru, small, rocky, rapid stream from Serra Jacobina between Bom Fim and Jacobina, about 11° S, Dorsal fin with a number of dark brown, 40°30' W, coll. J. D. Haseman, 8-XI-1907 (this specimen is mostly somewhat elongate dots, with a variable figured in Ellis, 1913, pi. XXVI fig. 3); four (paratypes), distribution over and membrane. Adipose rays FMNH 78361 [ex FMNH 54810, ex CM 3457] (two), ZMA fin usually with a small dark brown central or 113.595 [ex FMNH 54810] (one), CAS [IU] 13319 [ex CM

dorsal dot. Caudal fin with series of minute dark 3457] (one), si 23.3 to 30.8 mm, same data as holotype; three

FMNH 54808 [ex CM si about 21.0 three less (paratypes), 3455], to brown spots, forming up to more or about 33.4 mm, Brazil, Est. Bahia, Rio Agua Branca, coll. J. regular vertically or zigzaglike arranged narrow D. Haseman, 6-XI-1907; one (paratype), FMNH 54809 [ex lines. Anal fin usually with some faint spots at its CM 3456], si 33.7 mm, Brazil, Est. Bahia, Rio Zinga, coll. J. few of base; some specimens also have a such D. Haseman, 7-XI-1907.

Pelvic fins spots near the distal end of the rays. unpigmented. Dorsum of pectoral fins with light

Description. — brownish pigmentation, or with minute, irregular

into dark brown spots, sometimes arranged Morphometric and meristic data in table I and oblique lines. figs. 17 through 19. Dorsal and pectoral fin spines

distinctly shorter than adjacent rays. Snout in

Etymology. — front of nostrils naked. Two short mental barbels,

The specific name is derived from the Latin two pairs of long rictal barbels, reaching to gill from "fuscus" meaning dusky, dark, swarthy, and openings ventrally. Inner edge of pectoral fin fontanel the Latin "guttatus" meaning dappled, speckled, spine serrated (as in fig. 15-1). Anterior spotted, in allusion to the colour pattern. elongate oval (fig. 16k). Skin of intercoracoid

area naked.

Aspidoras maculosus n. sp. also (figs. 11, 15-1, 16k, 17-19,21) Colour in alcohol (fig. 11; see Ellis, 1913, pi.

XXVI fig. 3). — Ground colour of body and head

with several isolated roundish Aspidoras rochai [non R. von Ihering, 1907]; Ellis, 1913: pale tan. Body or

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maculosus Fig. 11. Aspidoras n. sp. Holotype.

Brazil, Est. Ceara, Poco Redondo, Rio dos CacAs at Crato, somewhat irregular brown spots, those on and Rio coll. R. S. de XII-1945. of Salgado system, Menezes, near midline longitudinal lateral body scutes

somewhat larger than the others. In some speci-

the tend be into — mens spots to arranged irregular Description.

horizontal rows. Morphometric and meristic data in table I and

Dorsal fin with faint, irregular or elongate figs. 17 through 19. Pectoral and presumably also

brown markings. Adipose fin. membrane with or dorsal fin spines distinctly shorter than adjacent

without minute brown Snout in of nostrils naked. Two short a spot. Caudal fin with up rays. front

mental of to four narrow vertical lines of small brown spots barbels, two pairs long rictal barbels,

on the rays. Anal fin presumably unpigmented reaching to or just beyond gill openings ventrally.

(the specimens are preserved during 69 years). Inner edge of pectoral fin spine moderately

Pelvic fins without pigmentation. Pectoral fin serrated (figs. 15 m-n). Anterior fontanel elon-

brown sometimes with some pigment on the gate oval (fig. 16-1). Skin of intercoracoid area

dorsum. the colour of the naked in Generally, pattern juveniles, with a few isolated minute

body and head of Aspidoras maculosus resembles ossifications in adults.

that of A. raimundi.

Colour in alcohol (fig. 12). — Ground colour of

Etymology. — body and head pale tan. Dorsal and caudal fins

in The specific name is derived from the Latin damaged most of the specimens. Dorsum of "maculosus" meaning spotted, dappled, pied, in head, snout, and opercles finely marbled with

allusion to the colour pattern. brown. Colour pattern on sides of body variable,

consisting of medium-sized to rather large ir-

regular dark brown blotches, most prominent

midlateral Aspidoras menezesi n. sp. along longitudinal body scutes, poste-

sometimes (figs. 12, 15 m-n, 16-1, 17-19,21) riorly united to form one elongate

horizontal blotch. Upper half of dorsolateral

Material examined. — One (holotype), UMMZ 147336, si body scutes with some ill-defined brown blotch-

41.8 mm, Brazil, Est. Ceara, Rio Granjeiro at Crato, 07°10' S, es, marbled, or cloudy. Lower half of ventro- 39°25' W, tributary on left bank of Rio Salgado, coll. R. S. de lateral body scutes unpigmented, except for a Menezes, 1945; twenty-six (paratypes), UMMZ 195951 brown dot connected with the on (twenty-one), ZMA 113.596 (five), si 20.3 to 34.0 mm, same pigmentation the midlateral in data as holotype; one (paratype), UMMZ 147340, si 21.9 mm, body scutes some specimens.

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Fig. 12. Aspidoras menezesi n. sp. Holotype.

Head, snout, and opercles finely marbled with Aspidoras poecilus n. sp. brown. (figs. 13, 15-o, 16m, 17-19,21)

Dorsal fin with without brown or some spots;

Corydoras cochui [non Myers & Weitzman, 1954]; Axelrod there are specimens with some brownish pig- et al., 1967: F-203.00 (colour photograph; popular ac- mentation in upper third of the fin. Adipose fin count). membrane unpigmented or with a faint brownish Corydoras rochai [non R. von Ihering, 1907]; Knaack, ! 970: Anterior of caudal fins spot. part damaged may CXXXI 333, pi. (photograph; popular account; discus-

bear one or two vertical or oblique rows of sion).

minute dark brown spots forming narrow lines.

Anal, pelvic, and pectoral fins unpigmented. Material examined. — One (holotype), IRScNB 560, si 29.2

mm, Brazil, Est. Mato Grosso, creek upstream of village

left bank of Rio Rio coll. P. Porori, Xingu, upper Xingu, J. Etymology. — Gosse, 31-X-1964; two (paratypes), USNM 191624 (one), This species is named in honour of Dr. Rui ZMA 113.597 (one), si 31.7 and 35.3 mm, largest probably a Simoes de who collected the Menezes, holotype male, Brazil, Est. Goids, Rio Araguaianear Aruana, 14°58' S, and paratypes. 51 °04' W, coll. H. R. Axelrod, 1960.

13. Fig. Aspidoraspoecilus n. sp. Holotype.

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Material examined. — Sixteen, MNRJ 997, si 15.0 to 24.6 Description. — coll. R. mm, Brazil, Est. Goias, Rio Palmas, Pfrimer, 1957; Morphometric and meristic data in table I and four, CAS 16015 [ex IU], si 28.9 to 30.3 mm, Brazil, Est. figs. 17 through 19. Dorsal and pectoral fin spines Goias, Corrego at Peixe into Rio Tocantins, 12°02' S, 48°36'

distinctly shorter than adjacent rays. Snout in W, coll. C. Ternetz, 31-1-1924; seventeen, CAS 16013 [ex IU],

front of nostrils naked. Two short mental barbels, si 23.7 to 31.4 mm, Brazil, Est. Goias, Rio Tocantins system, Vereda do Agostinho into Rio Cannabrava, coll. C. Ternetz, two pairs of rictal barbels, the longest reaching to 14-1-1924; one, CAS 16014 [ex IU], si 33.0 mm, Brazil, Est. gill openings ventrally. Inner edge of pectoral fin Goias, Rio Tocantins system, brook into Corrego da Porteira serrated Anterior spine moderately (fig. 15-o). into Rio Cannabrava into Rio Tocantins, coll. C. Ternetz, 15- fontanel round Skin of intercoracoid (fig. 16m). 1-1924; five, CAS 16012 [ex IU], si 26.9 to 33.2 mm, Brazil,

Sao coll. C. area with numerous isolated minute ossifications. Est. Goias, Corrego Domingo, Ternetz, 2-1-1924;

16011 si 27.7 Est. one, CAS [ex IU], mm, Brazil, Goias,

Colour in alcohol (fig. 13). — Ground colour of Corrego da Mula into Tocantins below mouth of Rio

about 14° 48°15' coll. C. body and head pale tan. Dorsum and sides of Maranhao, S, W, Ternetz, 1-1-1924; CAS 16009 [ex SU], si about 24.8 Brazil, Est. Goias, head, opercles, and snout marbled with tan and one, mm, Ribeirao Macaco Rio Tocantins system, Corrego Gomez into brown. Lateral sides of body with several prom- into Rio Maranhao, coll. C. Ternetz, 30-XII-1923. inent and weak blotches (see fig. 13, of left side

Description. — of holotype), which are slightly differenton both Morphometric and meristic data in table II. sides of the holotype. Dorsal and pectoral fin spines distinctly shorter Dorsal fin base with a large, oval, brown blotch from fourth and smaller than adjacent rays. Snout in front of nostrils spine through ray, a naked. Two short mental of brown blotch base of sixth and seventh barbels, two pairs on ray.

rictal to or almost to Both blotches extend somewhat over the dorsum long barbels, reaching gill

openings ventrally in some and to or of the body, the smaller one being connected specimens, base of fin in other with a dark solid blotch. Dorsal to the two basal slightly beyond pectoral spine specimens. Inner edge of fin (1) blotches there is a rather broad, horizontal pectoral spine finely serrated in MNRJ 997, (2) finely to mod- whitish stripe, extending to halfway the spine and erately serrated in CAS 16013 (fig. 15p), (3) followed by a large brown blotch. This blotch

the moderately serrated in CAS 16015 and CAS consists of a weakly pigmented part on to serrated in CAS membrane, alternated by dense pigmentation on 16014, (4) moderately strongly 16012, CAS 16011, and CAS 16009. Anterior the rays, forming vertical lines. Above this

fontanel (1) roundish, elongate oval, or slightly blotch, the dorsal fin is devoid of pigment. rhomboidal in the specimens in MNRJ 997, (2) Adipose fin spine with some scattered brown

roundish or pear-shaped in CAS 16015, (3) chromatophores along the base and near the elongate oval in CAS 16014, (4) elongate oval in middle; upper part of its membrane weakly most of the specimens in CAS 16013, though pigmented with brown. Caudal fin rays with

some have a broad oval anterior elongate brown spots, arranged into three or four fontanel, (5) broad oval in four specimens in CAS 16012, irregular, though somewhat vertical broad bars.

rather elongate oval in one specimen, slightly Anal fin with a series of minute brown spots (6) rhomboidal in CAS 16011 and CAS 16009. Skin arranged into an oblique U-shaped line. Pelvic

of intercoracoid area with isolated minute ossifi- fins unpigmented. Dorsum of pectoral fin spine

cations, which are more numerous in adults than and rays with scattered minute brown spots. in juveniles; in the latter they be even still Unossified ventral area whitish. Upper rictal may barbels with some brown pigment. undeveloped.

Colour in alcohol. — Most of the in Etymology. — specimens MNRJ 997 but of them The specific name is derived from the Greek are badly faded, some show four brown solid blotches the noiKtkos meaning variecoloured, pied, mottled, up to light on

in allusion the sides of the body. A blotch behind the base spotted, to colour pattern. just

of the last dorsal fin ray is the largest and covers

aff. Aspidoras sp. poecilus the greater part of the dorso- and ventrolateral

15p, 21) No is visible of (figs. body scutes. pigmentation on any

the fins. Most caudal fin are Aspidoras rochai [non R. von Ihering, 1907]; P. de Miranda rays, however,

MNRJ off Ribeiro, 1959: 8 (listed; in part; Rio Palmas, 997). broken near the base.

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meris- Table II. Morphometric characters expressed as ratios in sl (bd through hl), or as ratios in hl (sn through dcp), and some tic data of 17 specimens ofAspidoras sp. aff.poecilus.

Ids hi lbo wi D A C dbs vbs CAS no. si bd b.w lps sn ca dcp P, P, pas

— 25 23 4 16015 30.3 3.5 3.7 7.2 5.8 3.8 2.0 3.6 2.2 1 8

4 16015 30.0 3.4 3.8 7.1 6.0 3.8 2.0 3.8 2.3 — 1 9 25 22

4 16015 29.2 3.8 4.2 7.3 5.5 3.9 2.1 3.5 2.2 — 1 8 25 22

16015 28.9 3.3 3.8 8.0 5.7 3.6 2.0 3.8 2.4 — 1 9 25 23 3

16013 31.4 3.7 4.3 7.5 6.2 3.8 2.1 4.1 2.4 2.4 1 8 I,7,i i,5 ii,5 ,8 7/7 25 22 4

22 4 16013 30.7 3.7 4.2 6.3 6.5 4.0 1.9 4.2 2.1 1.9 1 8 I,7,i i,5 ii,5 ,8 7/7 25

16013 30.3 3.7 4.2 7.6 6.3 3.9 2.0 4.1 2.2 2.2 1 8 I,7,i i,5 ii,5 ,8 7/7 26 23 4

16013 29.7 3.6 4.0 7.1 6.0 3.7 2.0 4.8 2.4 2.1 1 8 I,7,i i,5 ii,5 ,8 7/7 25 22 4

16013 29.4 3.7 4.3 7.2 6.1 3.9 2.0 4.2 2.2 2.5 1 8 1,74 i,5 ii,5 ,8 7/7 25 22 4

16013 28.4 3.7 4.2 8.9 6.9 3.8 2.0 4.2 2.1 2.3 1 8 I,7,i, i,5 ii,5 ,8 7/7 25 22 5

5 16013 27.6 4.0 4.6 8.6 6.0 4.3 2.0 3.8 2.1 2.6 1 7 [,74 i,5 ii,5 ,8 7/7 26 23

22 16013 27.3 3.7 4.4 7.6 6.2 3.9 2.0 4.1 2.1 2.1 1 8 1.74 i,5 ii,5 .8 7/7 25 6

16014 33.0 3.6 4.0 7.7 5.8 3.8 1.9 3.9 2.4 2.1 1 8 25 22 4

16012 33.2 3.6 3.9 8.3 6.1 3.5 2.0 4.7 2.4 — 1 9 26 23 4

6 16012 31.3 3.5 3.8 8.0 5.9 3.4 1.9 4.7 2.4 — 1 9 25 23

16012 28.6 3.9 4.2 6.5 4.8 3.8 2.0 4.2 2.1 — 1 9 27 24 5

4 16012 27.3 3.8 4.1 7.0 5.4 3.6 2.0 3.9 2.3 — 1 8 27 24

A = anal fin; C = caudal fin; D = dorsal fin ;P,- pelvic fin; Pj = pectoral fin.

bad of fin is The specimens in CAS 16015, 16013, 16014, present condition their dorsal not

ideal for observation. The snout is marbled and 16011 all show a very similar colour pattern, finely after 50-odd of The snout is with brown in some The sides of the years preservation. specimens. somewhat marbled with light brown in some body have dark brown irregular dots, fewer and specimens. In other specimens the dorsal fin has larger and better defined than in the other CAS

The of the dots some light brown dots; sometimes the dorsal fin specimens here discussed. shape

is variable found in marble. Those the is plain. The caudal fin has up to three light as as along brown vertical or slightly oblique lines, which are meeting of dorso- and ventrolateral body scutes somewhat heavier in the lower lobe than in the are the largest. In the caudal fin there are up to

three vertical upper lobe. All specimens have a large number of narrow light brown or slightly

lines. irregular spots and dots on the sides of the body. oblique

Those along or near the meeting of dorso- and

ventrolateral body scutes are usually somewhat Aspidoras spilotus n. sp. larger than the remaining markings, and often (figs. 14, 15q, 16n, 17-19,21) tending to form a row. Again there is a great individual variability in minor details of the rochai Aspidoras [non R. von P. de Miranda colour the Ihering, 1907]; pattern. In some specimens spots on Ribeiro, 1959: 8 (listed; in part; MNRJ 8688, Cachoeira the sides of the body are isolated, sometimes they do Gusmao, Municipio de Ipu). are connected with other spots to form irreg-

Material examined. — ularly shaped blotches. One (holotype), ZMA 113.590,si 34.0 mm, Brazil, Est. Ceara, Riacho dos Macacos, tributary of Rio The specimens in CAS 16012 and 16009 are Acardu, coll. J. L. de Meneses, 10-XI-1969; twenty-nine still surprisingly well-pigmented (they were col- (paratypes), ZMA 112.284 (twenty-one), USNM 213568 (two), lected in the same period as the other CAS CAS 31834 (one), DNOCS 18112 (five), same data as The in specimens just mentioned). dorsal fin rays holotype; one hundred and forty-four (paratypes), MNRJ

8688 (one hundred and one of these specimens have dark brown spots forty-two), ZMA 113.591 (two), si 17.7 30.0 Est. Cachoeira do less horizontal to mm, Brazil, Ceard, Gusmao, arranged into three more or rows, Municipio de Ipu, coll. J. F. Cruz, 24-VIII-1952. one along the base, the second near the distal end of the and the third in between. rays, Basically, the other specimens seem to have the same Description. —

colour pattern of the dorsal fin. However, the Morphometric and meristic data in table I and

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14. ZMA sl 28.9 Fig. Aspidoras spilotus n. sp. Paratype, 112.284, mm.

base of last anal fin and between figs. 17 through 19. Dorsal and pectoral fin spines near ray, one

shorter than base of last anal fin and base of caudal fin. distinctly adjacent rays. Snout in ray

front of nostrils naked. Two short mental barbels, Dorsal fin with small, dark brown spots, mostly

of rictal in the half of the sometimes two pairs long barbels, reaching some- upper fin, forming times up to gill openings ventrally, but usually one or two narrow horizontal or slightly oblique shorter. Inner edge of pectoral fin spine finely to lines, irregularly arranged in other specimens,

blotch. The moderately serrated (fig. 15q). Anterior fontanel sometimes forming a small triangular

fin pear-shaped or oval (fig. 16n). Skin of inter- pigmentation in the dorsal is not restricted to

coracoid naked in with the it is much denser the area juveniles, minute rays only, although on isolated ossifications in than the membrane. Some adults, more numerous rays on pigmentation

than be at base of first and last two dorsal anteriorly posteriorly. may present additional fin rays. In some specimens there is an Colour in alcohol (fig. 14). — Ground colour of small, ill-defined blotch at base of middle dorsal body and head pale tan. Dense, dark brown fin base. Adipose fin spine and dorsal membrane pigment on dorsum and sides of head and snout, pigmented with brown. Caudal fin with to four for up and on opercles and cleithrum, except a rather broad, dark brown oblique or vertical in the sub- narrow pale tan curved line naked bars, better defined in some specimens than in orbital/preoccipital area. Snout sometimes slight- others; one of the paratypes has no bars but an ly marbled; the preoccipital/interorbital area Anal even, densely brown pigmented caudal fin. sometimes with lighter and darker brown round- fin sometimes plain, mostly with a faint brown ish spots. Body with prominent dark brown, oval dot of middle in the near the base the rays or spots along the meeting of dorso- and ventro- middleof these Pelvic fins without rays. pigment. lateral body scutes. There are four more or less Dorsum of pectoral fins usually with numerous the other isolated spots in one paratype. In minute brown with spots, or evenly pigmented specimens the spots are united, sometimes form- scattered brown. Belly whitish to yellowish. ing a single longitudinal bar from cleithrum to The pattern of the paratypes in MNRJ 8688 base of caudal fin. Most of the specimens are and ZMA 113.591 is faded. The midlateral blotch- intermediatebetween these two patterns. Dorsal es are the most prominent markings, especially to the midlateral series of spots the body may the centres of the blotches have remained, so have a light horizontal area. Other specimens are that most of the specimens do not show a in this The pigmented with light brown area. longitudinal bar. with dorsum of the body is light brown some

below last dorsal fin dense spots, usually just ray and just below adipose fin spine. Ventrum of

body (ventral to midlateral dots/bar) pale tan, but Etymology. —

there The usually are three dusky, ill-defined con- specific name is derived from the Greek

dark centrations of brown pigment, one in front anikazos meaning spotted, stained, in allusion

anal fin the colour of "spine" (unbranched ray), one at or to pattern.

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Fig. 15. Profiles of pectoral spines of a) Aspidoras rochai, lectotype, right spine, b) A. rochai,

paralectotype, left spine, c) A. raimundi, lectotype, left spine, d) A. lakoi, holotype, left spine, e)

A. pauciradiatus, paratype, left spine, f) A. albater, holotype, right spine, g) A. brunneus, holotype,

right spine, h) A. carvalhoi, holotype, right spine, i) A. eurycephalus, holotype, left spine, k) A.

fuscoguttatus, holotype, right spine, l) A. maculosus, holotype, right spine, m) A. menezesi,

holotype,left spine, n) A. menezesi, paratype, ZMA 113.596,sl 30.7 mm, left spine, o) A.poecilus,

aff. CAS sl 30.3 left and holotype, right spine, p) A. sp. poecilus, 16013, mm, spine, q) A. spilotus, holotype,right spine.

earliest DISCUSSION Pliocene time was transformed into the

Rio Amazonas, flowing to the Atlantic Ocean by

and the intensive of the As shown in figs. 20 21, the genus Aspidoras uplift Andean Cordillera in has rather restricted from the & a distribution, Pliocene time (Damuth Kumar, 1975: 874).

course of the Rio in the West to the this area and be upper Xingu Corydoras occupied might con- sidered upper course of the Rio Itapicuru in the East, more advanced than Aspidoras.

in the North and from the Rio Acardu system to Aspidoras rochai, A. raimundi, A. lakoi. A. the of the Rio in the South. upper course Parana pauciradiatus. A. albater, A. brunneus,A. carvalhoi,

The absence of Aspidoras in the rest of the A. eurycephalus, A. fuscoguttatus. A. maculosus, A. continent, particularly its absence in the main menezesi, two forms of Aspidoras sp. aff. poecilus,

Rio Amazonas basin, is of zoogeographical in- and A. spilotus were each collected in such

this restricted that terest. Aspidoras apparently never occupied areas endemism may be suspected area, which consisted of immense lakes in middle for most of the species. In fact, only A. poecilus

Miocene time, and which in late Miocene or has been found in two widely scattered places. As

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Fig. 16. Anteriodorsal view, showing fontanels and nuchal plates in a) Aspidoras rochai,

lectotype, b) A. raimundi, lectotype, c) A. lakoi, holotype, d) A. pauciradiatus, ZMA 113.586, sl

20.1 mm, e) A. albater, holotype, f) A. brunneus, holotype, g) A. carvalhoi, holotype, n) A.

eurycephalus, holotype, i) A. fuscoguttatus, holotype, k) A. maculosus, holotype, l) A. menezesi,

holotype, m) A.poecilus, holotype, n) A. spilotus, holotype, and o) Corydoraspygmaeus, holotype.

in the immediate For- far as we know, no Aspidoras specimens are course neighbourhood of mixed with ofthe in taleza. Three other were collected from up members genus Corydoras species museum collections, as we have been able to rivers about Fortaleza: A. carvalhoi, A. menezesi, examine the majority of the representatives of and A. spilotus. The latter species, of which 169

be the latter genus. specimens are available, seems to closely

Aspidoras rochai (type-species of the genus) is related to A. rochai. Especially the colour pat- known only from two badly preserved specimens terns of A. rochai and A. spilotus are quite similar,

from is fact that the colour A. originating Fortaleza. There no exact despite the pattern of

from col- is known R. indication which locality they were rochai only from von Ihering's

of lected; there are several rivers with an undefined original description. The two specimens A.

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Fig. 17. Variation in ratios of body depth (bd), body width (bw), dorsal fin spine length (lds), and

pectoral fin spine length (lps) in standard length (sl) in the species ofAspidoras. Verticals indicate

primary type-specimens.

rochai almost the of are now completely depigmented, near type locality A. rochai, A. carvalhoi

the fins are damaged, and even R. von Ihering and A. menezesi show such distinctly different

already was unable to make an accurate descrip- characters that they cannot be confused with A.

tion of the colour pattern. The two specimens of rochai.

A. rochai measure 39.9 and 37.8 in whereas raimundi from the mm si, Aspidoras upper Rio Par-

the largest specimen ofA. spilotus has a si of 34.0 naiba is known from 63 specimens, all collected

from the It described in the mm, which may account for some difference in type-locality. was

A. and A. A. but proportions. However, rochai spilotus same year as rochai, not as an Aspidoras as

differ in the following characters: length bony a Corydoras. Until now, A. raimundi was never

orbit 5.3 to 5.4 in A. rochai against 3.7 to 5.0 in A. figured in literature. A. raimundi seems most spilotus; body width 3.8 in A. rochai against 3.9 to closely related to A. menezesi. Several morpho-

4.3 in A. spilotus; 6 pre-adipose scutes in A. rochai metry and meristic characters are alike. A.

raimundi against 3 to 4 in A. spilotus. Moreover, A. rochai is a smaller species (si up to 32.6 mm tends have 41.8 in to a greater body depth (3.4 to 3.8 against up to mm A. menezesi). A. menezesi against 3.7 to 4.3 in A. spilotus), shorter dorsal fin never shows a dark, prominent blotch in the

6.4 9.0 in A. and and spines (9.0 against to spilotus), dorsal fin, tends to have a somewhat longer shorter pectoral fin spines (6.2 to 7.8 against 5.4 snout (1.8 to 2.0 against 1.9 to 2.1 in A. raimundi),

6.3 to in A. spilotus). Both specimens of A. rochai and a narrower interorbital (2.2 to 2.7 against 2.0 have from the 2.3 in a habitus distinctly different 169 to A. raimundi). Although we regard A. specimens of A. spilotus, the former species being raimundiand A. menezesi distinct at specific level more robust. The two other species occurring (based on the material available), they might

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Fig. 18. Variation in ratios of head length (hl) in standard length (sl), and snout length (sn), length bony orbit (lbo), and interorbital width (wi) in head length in the species of Aspidoras.

Verticals indicate primary type-specimens.

when from all other species of the its low prove to be conspecific specimens genus Aspidoras by number of dorso- and ventrolateral more localities in the area become available. body scutes

in the other Aspidoras lakoi, the second species described in (23/20 against 25/22 or more species),

based on the its low number of dorsal fin (6 against 7 in Aspidoras, was single holotype. rays

series of 23 collected in the other its short snout Fortunately, a topotypes; species), extremely (2.5

available for 2.1 or less in the other and its the same year as the holotype, was against species), by

examination. A. lakoi is known from the Rio colour pattern. A. pauciradiatus has a more

serrated fin Parana system, from a tributary of the Rio strongly pectoral spine than its

Grande. In the same river system A. fuscoguttatus congeners. differences Of the describe occurs. Apart from in colour pattern, species we as new, three are

from in the viz.: A. fuscoguttatus differs A. lakoi easily recognizable by their colour pattern,

characters: dorsal fin A. albaterwith broad solid following a longer spine oblique brown bars, A.

8.7 8.7 brunneus half the solid (6.7 to against to 10.4 in A. lakoi); a longer with the upper of body snout (1.7 to 1.9 against 1.9 to 2.0 in A. lakoi); and dark brown, and A. carvalhoi with hardly any

less pre-adipose scutes (3 to 5 against 6t013 in A. pigmentation.

Besides lakoi). Moreover, A. fuscoguttatus tends to have a Aspidoras albater, two other species

orbit in tributaries of the larger bony (5.7 to 6.6 against 6.5 to 8.0 in occur upper Rio Tocantins:

a aff. and A. lakoi), and narrower interorbital (2.1 to 2.4 Aspidoras sp. poecilus, A. eurycephalus. In against 2.0 to 2.2 in A. lakoi). the upper course of the Rio Araguaia two species

Aspidoras pauciradiatus (originally described in were found together near Aruana: A. poecilus is the genus Corydoras) easily distinguished from (two paratypes) and A. pauciradiatus (holotype

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Fig. 19. Variation in ratios of depth caudal peduncle (dcp) in head length (hl) and variation in

number of dorsolateral body scutes (dbs), ventrolateral body scutes (vbs), and pre-adipose

scutes (pas) in the species of Aspidoras. Verticals indicate primary type-specimens.

of the and paratype). The specimens here considered as species genus Aspidoras.

aff. all maculosus is known from three Aspidoras sp. poecilus are badly preserved Aspidoras near-

from tributaries of the Rio situated tributaries to the Rio specimens upper by upper Itapicuru.

Tocantins, which resemble A. poecilus in many It resembles A. menezesi most, but it has smaller

details. However, there are two forms: one with and better defined roundish dots rather than the

in A. the blotches on the body more conspicuous and irregularly shaped dots menezesi. Moreover,

larger than in A. poecilus, the other form with A. maculosus differs from A. menezesi in the comparatively smaller and less conspicuous following characters: shorter dorsal fin spines

markings than in A. poecilus. Both forms differ (7.5 to 9.0 against 6.2 to 7.2 in A. menezesi),

from each other and from A. poecilus in relative shorter pectoral fin spines (5.8 to 6.8 against 4.9

length of dorsal and pectoral fin spines. Further to 5.6 in A. menezesi). Furthermore, A. maculosus

differences but these be due to has 3to 4 5t06 in are found, may pre-adipose scutes against

the imperfect conditionof the material. menezesi; it also tends to have less body scutes

Aspidoras eurycephalus, also from the upper Rio (dbs/vbs 25-26/22-23 in A. maculosus against

in Tocantins, was collected almost simultaneously 26-27/23-24 A. menezesi). Finally, A. maculosus

of the of tends have less 4.0 with the greater part specimens Aspi- to a deep body (3.8 to against

doras also to 3.8 in sp. aff. poecilus. Although they are 3.3 A. menezesi), a more slender body

badly preserved specimens, their broad inter- (4.2 to 4.4 against 3.8 to 4.2 in A. menezesi), and a

orbital not only differs from A. poecilus and less deep caudal peduncle (2.0 to 2.2 against 1.8

2.0 in Aspidoras sp. aff. poecilus, but also from all other to A. menezesi).

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Figs. 20-21. Occurrence ofAspidoras species.

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