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http://www.jstor.org Copeia,1989(2), pp. 332-344

Pteroglanis manni Eigenmann and Pearson, a Juvenile of Sorubimichthysplaniceps (Agassiz), with a Review of the Nominal of Sorubimichthys (Pisces: )

JOHN G. LUNDBERG, PEDRO NASS AND FRANCISCO MAGO-LECCIA

Recently collected growth series of Sorubimichthys planiceps from the Apure and Orinoco rivers in the llanos of central Venezuela demonstrated marked ontogenetic change from juvenile to adult in coloration and fin and snout mor- phology. Comparison of the holotype and original description of Pteroglanis manni to the juveniles of S. planiceps revealed close similarity, suggesting that these taxa are conspecific. The nominal species of Sorubimichthys are reviewed based on examination of the literature, available types and other specimens. Sorubimichthys spatula (Agas- siz), the type species, and S. planiceps (Agassiz) are likely conspecific but are not synonymized due to lack of information on S. spatula whose holotype (the only specimen) is destroyed. A syntype of S. planiceps has been located in the MHNN, and is designated the lectotype. Platystoma artedii Giinther, long treated as a synonym of S. planiceps, is reallocated to the Pseudoplatystoma. Sorubim- ichthys ortoni Gill is confirmed as a junior subjective synonym of S. planiceps. The holotype of S. gigas (Giinther) is identified as belonging to the genus Brachy- platystoma.

Series de desarrollo de Sorubimichthys planiceps colectadas recientemente en los rios Apure y Orinoco en los llanos centrales de Venezuela revelan cambios ontogeneticos notables en la morfologia del hocico, aletas y la coloracion entre los ejemplares juveniles y adultos. La comparacion del holotipo y la descripcion original de Pteroglanis manni con los juveniles de S. planiceps presenta similar- idades que sugieren que estos taxa son coespecificos. Las especies nominales de Sorubimichthys fueron revisadas en base al examen de la literatura, tipos disponibles y otros ejemplares. Sorubimichthys spatula (Agassiz), la especie tipo, y S. planiceps (Agassiz) son aparentamente coespecificas pero no fueron sinonimizadas debido a la ausencia de informacion suficiente sobre spatula cuyo holotipo, el unico ejemplar, fue destruido. Un sintipo de S. planiceps pudo ser localizado en el MHNN, y fue designado el lectotipo. Platys- toma artedii Giinther, desde hace mucho tiempo considerado como un sinonimo de S. planiceps, fue asignado al genero Pseudoplatystoma. Sorubimichthys ortoni Gill fue confirmado como un sin6nimo subjectivo menor de S. planiceps. El holotipo de S. gigas (Giinther) fue identificado como un Brachyplatystoma.

THE family Pimelodidae presents one or a handful of individuals. These especially some of the more challenging problems remain isolated in our developing conception in South American ichthyology. This diverse of pimelodid classification, awaiting additional family contains over 300 nominal species and material or data pertaining to their relation- several more pending description. Despite ships and status. This paper identifies and at marked morphological diversification underly- least partly resolves a few such problems with ing the more than 50 nominal genera, it is only the nominal genera and species of Sorubimichthys recently that genera and higher groups have and Pteroglanis. The impetus for this work came begun to be characterized in a phylogenetic from the study of recently collected larval and framework (Stewart, 1986; Lundberg and juvenile pimelodids and emphasizes the impor- McDade, 1986; Lundberg et al., 1988). Many tance of developmental information in the sys- nominal pimelodid taxa are known from just tematics of South American fishes.

? 1989 by the American Society of Ichthyologists and Herpetologists LUNDBERG ET AL.-REVIEW OF SORUBIMICHTHYS 333

GENUSSORUBIMICHTHYS as, or assigned to, Sorubimichthyshas presented questions of one sort or another. We are able Agassiz (in Spix and Agassiz, 1829) estab- to resolve or offer additional information and lished the catfish and described genus Platystoma opinions on all of these. five member species, including P. planiceps and Whitehead and Myers (1971) reported that P. To these two nominal spatula. species, Spix, the type of planiceps was destroyed in Munich in of the same volume the plates (here repro- during World War II. However, M. Kottelat duced as 1 the Fig. A-B), simultaneously applied (pers. comm.) located a probable syntype of names Sorubim and Sorubim pirauaca jandia. planiceps in the MHNN, that closely matches Whitehead and the Myers (1971) analyzed Agassiz's original description and Spix's plate. of nomenclature of and problems Spix Agassiz's Photographs of this 458 mm specimen (fish "Brazilian Fishes" and concluded that Agassiz's lengths reported in this paper as SL) sent to us names and are valid. Bleeker planiceps spatula by J.-P. Haenni clearly show it to be the species name when he used (1862) accepted Spix's jan- that is everywhere called S. planiceps (see below). dia as the of his So- (spatula) type species genus The syntype that we designate as the lectotype did rubimichthys.Bleeker, however, not assign is labelled "Platystomaplaniceps Agassiz (Sorubim to or oth- pirauaca (planiceps) Sorubimichthys any pirauaca Spix), Bresil, 811." er genus. Gunther (1864) continued to recog- Platystoma artedii Gunther (1864) was based nize sev- Agassiz's Platystomacontaining, among on the figure of"Mystus No. 6," Figure 6, Plate eral others, both planiceps and spatula. 29, in volume 3 of Seba (Levrault 1827-1828). and ac- Eigenmann Eigenmann (1888, 1890) Study of this figure leads us to conclude that Bleeker's cepted Sorubimichthys,noted that the artedii is based on a specimen of Pseudoplaty- name is in the Platystoma preoccupied Diptera, stoma, probably P. fasciatum. Contrary to Gun- and used both Agassiz's names, allocating spat- ther's (1864) original description, the snout, ula and to A planiceps Sorubimichthys. summary while apparently a little longer than the lower of and Eigenmann Eigenmman's proposed syn- jaw, does not project greatly (not nearly as much for these is as follows: onymies species as in Sorubimichthysplaniceps). The figure shows other characteristics of artedii that both match Sorubimichthys spatula (Agassiz, in Spix and Pseudoplatystomaand clearly distinguish it from Agassiz, 1829) S. such as the cranial fontanelle Platystoma in and planiceps, deep spatula Agassiz, Spix Agas- that extends to the caudal siz, 1829 posterior eye, spotted fin, mental barbels that extend beyond the head Sorubimjandia Spix, in Spix and Agassiz, 1829 to the pelvic fins, and adipose fin inserted in Sorubimichthysplaniceps (Agassiz, in Spix and advance of the anal fin. Body color pattern is Agassiz, 1829) unfortunately not illustrated. Also of interest is Platystomaplaniceps Agassiz, in Spix and Agas- the caption of the figure that parenthetically siz, 1829 notes "Pimelodusfasciatus Lacep., Silurusfascia- Sorubim pirauaca Spix, in Spix and Agassiz, tus Gmel.," which we suppose refer to Pseudo- 1829 platystomafasciatus (Linnaeus). We recommend Platystoma artedii Gunther, 1864 that Platystomaartedii Gunther be removed from Sorubimichthysortoni Gill, 1870 synonymy of S. planiceps and be treated as a subjective junior synonym of Pseudoplatystoma Eigenmann and Eigenmann (1888, 1890) also fasciatus (Linnaeus). allocated Platystomagigas Gunther, 1872, to So- Sorubimichthysortoni Gill (1870) is based on a rubimichthysmaking a total of three nominal single specimen from the Peruvian Amazon species in the genus. housed at the USNM. Following examination Miranda Ribeiro (1911) reported specimens of this specimen we concur with Eigenmann and that seemed to share the supposed diagnostic Eigenmann (1888, 1890) that it is S. planiceps. characters of these nominal species and syn- onymized the three under the name planiceps. The type of Platystoma gigas Gunther (1872) Miranda Ribeiro's lead has not been followed is a stuffed, mounted and apparently varnished, and the Eigenmann and Eigenmann (1888, fish about 2 m TL from the Rio Huallaga in the 1890) synonymy persists in the latest catalogues Amazon Basin. Information on this specimen of the Pimelodidae (Gosline, 1945; Fowler, and photographs were provided by G. Howes 1951). Each of the nominal species described and Mo Tian-pei of the BMNH. Based on these 334 COPEIA, 1989, NO. 2

I"

_-

I I M 1

W W_ .

A

.-- ^ ^ . : ..

B

Fig. 1. A) Sorubimichthysplaniceps (Agassiz), Sorubim pirauaca of Spix, 1829, pl. 12; B) Sorubimichthysspatula (Agassiz),Sorubim jandia of Spix, 1829 pl. 14. After Spix and Agassiz (1829). sources gigas is at once distinguished from S. Plate 14 of jandia (spatula) suggest a fish gen- planiceps by differently shaped palatal teeth (Fig. erally similar to S. planiceps (Fig. 1A-B). The 2A, E), heavy and recurved jaw teeth, approx- head of S. spatula is much like that of S. planiceps imately equal anal-fin and adipose-fin bases (Fig. in snout projection and outline, presence of small 3), and a relatively deep body (ca 2.3 in pre- black spots, and absence of deep cranial fon- dorsal length vs 3.8 in S. planiceps). These same tanelles (shown clearly to be present in adjacent characters plus the projecting upper jaw and illustrations of species of Pseudoplatystoma).The flattened head are sufficient to identify gigas as spotted color patterns of the dorsal, adipose and a specimen of the distinctive species Brachypla- paired fins are also similar to those of S. plani- tystomafilamentosum (Lichtenstein). ceps. Other similarities with S. planiceps are the Finally, the status of S. spatula is the most relatively short mental barbels and subfalcate difficult problem in the of the Soru- form of the dorsal fin. Based on these shared, bimichthys.Like gigas, S. spatula was described mostly unique similarities we suspect that S. from a single large holotype, in this case about spatula and S. planiceps represent the same 1 m. Like S. planiceps, the type of S. spatula was species. Nevertheless, the illustration of S. spat- destroyed during World War II, but unlike S. ula shows differences from S. planiceps. These planiceps, no syntypes exist. Furthermore, we are the apparent lack of the distinctively bold have encountered no subsequent material re- color pattern of the body, the presence of ser- ferred to S. spatula in the literature or in col- rations on the anterior margin of the pectoral lections. The large sorubimine pimelodids gen- spine, the insertion of the anal fin behind the erally have widespread distributions and are adipose-fin origin, and the lyriform upper and common in commercial catches. It is, therefore, lower lobes of the caudal fin. unlikely that S. spatula represents a species that Thus, the original and only source of infor- was collected just once, but its identity is prob- mation on S. spatula indicates a fish obviously lematical. close to S. planiceps, but with some apparent Agassiz's original description and Spix's color differences. It is possible that the differences LUNDBERG ET AL.-REVIEW OF SORUBIMICHTHYS 335

B and rare literature sources accounted for far A more than 50% of work time in this study. Ich- thyologists contemplating research on the large South American , that often were named, yet insufficiently described, more than once, should be prepared for considerable cor- respondence, travel and, in some cases, incon- clusively resolved taxonomies. I I Sorubimichthysplaniceps Sorubimichthysplaniceps is a large, commer- cially important species that is common in the C D major rivers of the Orinoco and Amazon basins. This species is among the most distinctive of the "shovel-nosed" pimelodid catfishes that Schultz (1944) united in the subfamily Sorubiminae (more or less the equivalent of Bleeker's So- rubimes, 1862). Whereas we do not endorse an hypothesis of monophyly for this subfamily, it I \ is apparent that the closest relatives of S. plan- iceps are included in it. In particular, we have evidence pointing to a monophyletic group in- E cluding Sorubimichthys,Pseudoplatystoma and So- rubim (Lundberg and Nass, unpubl.). In this study, therefore, we focus our comparisons on the sorubimines. The adults of Sorubimichthys planiceps are differentiated from the species of Brachyplatystoma(except B. filamentosum), Gos- linia, Phractocephalus, , Hemisoru- Fig. 2. Silhouettes of upperjaw dentitions:A) So- bim, , Paulicea, Perrunichthys,Platynema- Sorubimlima; rubimichthysplaniceps; B) C) Platystoma- tichthys and Steindachneridion by the extreme tichthyssturio; D) E) Pseudoplatystomafasciatum;Brachy- prolongation of the upper jaw (premaxillae) to platystomafilamentosum. approximately one third of the snout length. Sorubimichthysplaniceps is distinguished from all are due to preservation artifacts or illustration sorubimines (i.e., the foregoing genera plus B. errors, but such cannot be demonstrated. Al- filamentosum, Pseudoplatystoma,Platystomatichthys though we strongly suspect that S. spatula is and Sorubim) by the form of the tooth patches conspecific with S. planiceps, and not a large- on the upper jaw and palate (Fig. 2). Further, bodied species that has escaped capture and no- Sorubimichthysplaniceps has a unique color pat- tice since the early 19th century, this cannot be tern (Figs. 3D, 4D): an ash-gray dorsum with unequivocally demonstrated with available in- small darker spots; sides with a white band, bor- formation. Reluctantly we recommend that S. dered below by a gray or brown band that may spatula and S. planiceps be treated as valid, with be broken in blotches; venter white usually with the former being questionable as a separate bi- some large spots; dorsal and adipose fins spot- ological entity. This is an unfortunate result ted. We have examined specimens from because it likely inflates artificially the species throughout the Orinoco (Venezuela, Colombia) level diversity of these important and conspic- and Amazon (Brazil, Ecuador, Peru, Bolivia). uous catfishes. A bolder approach is to treat Although individual coloration is variable in the these as conspecifics with S. planiceps the senior density of spotting and the development of the synonym, following Miranda Ribeiro (1911) as ventrolateral band, there is no detectable pat- first reviser. tern of geographic or other taxonomically sig- In closing this section of the paper we note nificant variation in color, counts or form. A that the job of gathering data on the nominal single widespread species is represented. species of Sorubimichthysfrom type specimens The juveniles of S. planiceps are now known 336 COPEIA, 1989, NO. 2

A

.I 15 r

*" "

B

c

Pe * -- ~ aj_ I_ D

Fig. 3. Growth series of Sorubimichthysplaniceps, left lateral views: A) 10 mm; B) 11 mm; C) 33 mm; D) 550 mm.

from a recently collected developmental series present. Upper and lowerjaws about equal. Nu- from the Rio Apure on the llanos of central merous irregularly scattered melanophores over Venezuela and a few specimens from elsewhere. entire head and body. They occur along river banks hovering in vege- tation and are easily captured with dip nets. Circa 11-15 mm (Figs. 3B, 4B).-Although these This material presents the following features: specimens are only slightly longer than the pre- ceding, they exhibit important ontogenetic Circa 10 mm(Figs. 3A, 4A).-Dorsal- and ventral- changes. Dorsal, adipose and pelvic fins present; fin folds present; pectoral fins small; caudal fin anal fin scarcely developed, ventral-fin fold still strongly "heterocercal" in form, with few prin- large. Pectoral fins broad and elongated, reach- cipal rays. Long maxillary and mental barbels ing to near pelvic fins; lower lobe of caudal fin LUNDBERG ET AL.-REVIEW OF SORUBIMICHTHYS 337

D

Fig. 4. Growth series of Sorubimichthysplaniceps, head in dorsal view: A) 10 mm; B) 11 mm (some pectoral rays concealed by body); C) 33 mm; D) 550 mm. developed. Upper jaw projecting a little beyond folds that nearly reach the adipose and anal fins mandibles. By ca 15 mm palatal dentition com- respectively. Anal fin developed and continued prised of few scattered teeth (Fig. 5A). Body, forward to urogenital papilla as a high fin fold. top and upper sides of head uniformly dark Pelvic fins broadly rounded, not elongated. Pec- brown. Cheeks pale. Dorsal, caudal and pector- toral fins broad and prolonged beyond origin al fins dark brown. of pelvic fins to level of anus. Pectoral spine short, slender and sharp, without dentations. Circa 21 mm.-This specimen is important be- Upperjaw projecting completely beyond lower, cause it is nearly identical in size and very similar exposing premaxillary bands of teeth. Palatal overall to the holotype of Pteroglanis manni (see dentition comprised of few scattered teeth. below). All fins differentiated. Dorsal fin short, Dorsal and paired fins very dark. Cheeks and dorsal spine scarcely developed. Adipose con- dorsum speckled with large melanophores. tinued forward as a low fin fold. Caudal deeply Margins of branchial membrane, anal fin, cau- forked, its lobes pointed but not elongated. Both dal fin and adipose fin hyaline. Maxillary barbels caudal fin lobes continued forward by low fin banded. 338 COPEIA, 1989, NO. 2

*A Ao

p ' 4 p '4 " * . 11- .-^V . .^ t . Al-

Fig. 6. Living juvenile of Sorubimicthysplaniceps, 40 mm, Rio Apure, Venezuela.

middle anal-fin rays, and the dorsal-fin margin and middle caudal rays become hyaline. At ca 113 mm the pectoral fins were still enlarged, extending beyond the pelvic-fin origins. At 113 mm the palatal dentition (Fig. 5B) is comprised of three pairs of tooth patches that subsequently fuse (Fig. 2A). The adult color pattern (Figs. 3D, 4D) and pectoral, pelvic and caudal fin shape appear by at least 200 mm (smallest fish observed by us with these characters). Early in their development the juveniles of S. are at once distinct from their adult 5. Silhouettes of dentitions of planiceps Fig. upper jaw ju- form as far as from the venile 15 mm, scale bar and, known, juveniles Sorubimichthysplaniceps: A) of all other From the Rio Orinoco 0.5 mm; B) 113 mm, scale bar 4 mm. pimelodids. Basin,juveniles of the following taxa were avail- able for comparison: Brachyplatystomafilamen- tosum, B. juruense, B. rousseauxi, B. vaillanti, Circa 33 mm (Figs. 3C, 4C).-All fins differen- Hemnisorubimplatyrhynchos, Leiarius marmoratus, tiated and lacking fin folds. Caudal lobes slender Megalonemnaplatycephalum, Megaloneina sp. (cf. and elongated. Pelvic fins broad and elongated. xanthum), Phractocephalus hemiliopterus, Pimelo- Pectoral fins rounded and greatly elongated, della spp., Pinelodus cf. altipinnis, P. blochi, P. reaching beyond the origin of pelvic fins. Pec- pictus, Pinirampus pirinampu, Pseudopimeloduscf. toral spine slender, sharp, with a few small den- apurensis, Pseudoplatystomafasciatum, Pseudopla- tations on posterior margin. Head much de- tystomatigrinum, Rhamdia sp., Sorubim lima and pressed; upper jaw markedly projecting beyond Sorubimcf. latirostris.These series present a sur- lower. Most of head, body and fins uniformly prisingly rich diversity of juvenile characters dark brown. Margin of branchial membrane, (Mago-Leccia et al., 1986) and none possess the anal fin and margin of adipose fin hyaline. singular coloration and the fin and head form of juvenile Sorubimichthysplaniceps. The juve- Circa 40-113 mm.--An individual measuring niles of Pseudoplatystomaare most similar to those about 20 mm captured on 30 June 1984 was of S. planiceps in having depressed heads, with raised in a laboratory aquarium for 6 mo where somewhat prolonged upper jaws and elongated it grew to 113 mm. At about 40 mm (Fig. 6) a pectoral fins. In contrast, however, Pseudopla- white band appeared on the sides, extending tystomajuveniles have pointed pectoral fins, not from the pectoral fin to the caudal peduncle. broadly rounded fins, their anal and adipose fins This band is retained throughout life (Fig. 3D). are not hyaline but have dark bands, and their Between 60 and 70 mm a series of small round- heads have a sharply delimited pattern of coun- ed spots appears on top of the head and nape. tershading instead of an almost uniformly dark Also, an elongate dark spot develops over the color. Sorubim, with markedly projecting upper LUNDBERG ET AL.-REVIEW OF SORUBIMICHTHYS 339

Fig. 7. Holotype of Pteroglanismanni. After Eigenmannand Pearson (1924).

jaws in juveniles and adults, lack the greatly a flying fish" (Eigenmann and Pearson, 1924). enlarged pectoral fins, and have a much elon- Further, Eigenmann and Pearson described gated lower caudal fin lobe, larger more lateral these fins as "very wide and long, extending eyes, and a distinctively different color pattern. almost to the anal." W. M. Mann, collector of the holotype, mentioned the pectoral fins in his field notes. The illustration of the Pteroglanis manni holotype (Fig. 7) shows that the left pectoral fin was then most- W. M. Mann, circa 1921 (in Eigenmann and ly broken, although a few of the posterior fin Pearson, 1924), on the capture of the holotype rays were still long enough to have reached the of Pteroglanis manni wrote: ". . . it looked as pelvic-fin base. Both pectoral fins of the holo- much like something else as it did like a fish." type (Fig. 8A) are now badly broken and their This taxon was described by Eigenmann and original extent cannot be confirmed. Despite Pearson (1924) based on a single specimen tak- this damage, there is no reason to doubt the en in Bolivia in Jan. 1922 (not 1921 as pub- veracity of the original description of the pec- lished) from the Rio Negro of the Rio Beni toral fins. This is a critically important feature drainage, Estado Beni, ca 67?W, 13?S. The 21 because, as noted above, such elongated and mm holotype of P. manni (CAS 59623, origi- broadened pectoral fins are known to occur only nally Indiana University 16001), is the only in S. planiceps (Figs. 3-5). The elongated pec- specimen known to us. According to R. M. Bai- torals of juveniles of Pseudoplatystomaspecies, ley (pers. comm.), Pteroglanis Fowler (1934a) is the only other pimelodid genus known to have ajunior homonym of Pteroglanis Eigenmann and such greatly elongated pectorals, are narrow Pearson (1924). Fowler (1934b) established and pointed. The right pectoral fin of the ho- Pteropsoglanis to replace his homonym. lotype of Pteroglanis is less damaged than its Our attention was drawn to the similarity be- antimere and contains 10 soft fin rays; this is tween P. manni and juvenilies of Sorubimichthys the modal soft pectoral ray number of S. plan- planiceps by Eigenmann and Pearson's (1924) iceps (cf. Pseudoplatystomaspp. with modally nine illustration reproduced here (Fig. 7; see also soft pectoral rays). Both pectorals of the holo- Fig. 8). Although the illustration is flawed in type are darkly pigmented as in S. planiceps. The certain respects, it shows obvious similarities to left pectoral spine (not clearly illustrated by Ei- S. planiceps, especially the enlarged pectoral fins, genmann and Pearson, 1924; Fig. 8A) is com- depressed form of the head, and pigmentation plete; it is slender (about equal in width to a pattern. Examination of the holotype of P. man- soft fin ray), pungent, without marginal dentic- ni (hereafter in this section called "the holo- ulations, and equal in length to the bony inter- type") confirms its close similarity to S. planiceps orbital width. In this the pectoral spine of the juveniles between 21 and 39 mm in the features holotype is essentially identical to that of the 21 mentioned in the original description and oth- mm specimen of S. planiceps. The right pectoral ers as reviewed below. Sorubimichthysplaniceps spine is broken distally. also occurs in the Rio Beni. The heads and mouths of the holotype and The primary character given in the diagnosis comparably sized juveniles of S. planiceps are of Pteroglanis is the great enlargement of the much broadened. In these wide-mouthed fishes pectoral fins "which give it the appearance of the gape is visible laterally where the mouth 340 COPEIA, 1989, NO. 2 angle and and fleshy rictal fold terminate below A the posterior nostril and eye. Both the original illustration (Fig. 7) and description of P. manni are incorrect in suggesting that the mouth is terminal. Due to poor preservation and distor- tion, the snout and upper lip of the holotype are creased and folded ventrad. When carefully straightened, the snout and upper jaw protrude well beyond the chin, such that the premaxillary tooth bands far overhang the mandibular teeth (Fig. 8B). This form of the mouth is like that exhibited by S. planiceps adults and juveniles 21 mm and greater. These points of similarity are reflected in length relationships of the head and snout (Fig. 9). The relative lengths of the maxillary and mental barbels are similar in the and holotype Fig. 8. Head and anterior body of holotype of the maxillaries extend to about the S. planiceps: Pteroglanismanni, dorsal and ventral views. base of the pelvic fin, the outer mentals reach to the pectorals, exceeding the inner mentals by 2-4 times. Our 33 mm specimen of S. plan- head as measured from the tip of the snout. iceps (Fig. 3C) has abnormally short outer men- This character is better seen in dorsal views tal barbels. (Figs. 4, 8) than lateral view because of the ap- The caudal fin of the holotype is broken prox- parent foreshortening of the broad snout in imal to the fork. Although this is the condition perspective illustrations. Eigenmann and Pear- of the fin originally illustrated (Fig. 7), Eigen- son (1924) wrote incorrectly that the orbit lacks mann and Pearson (1924) stated that both lobes a free margin in P. manni. In the holotype, as of the caudal fin were very long. This is another in S. planiceps, there is a deep orbital furrow point of distinctive resemblance between the below, in front of and behind the eye. Above holotype and juveniles of S. planiceps (Figs. 3, the eye, however, the orbital rim is not folded 5). inward but is actually slightly raised. The taxo- The dorsal fin of P. manni, S. planiceps and nomic distribution of this feature among pi- that of most other pimelodids contains a spine melodids is not yet determined although it is and six soft rays. The spine and fin of the ho- known that several species have completely free lotype are now mostly broken; enough of the orbital rims whereas others lack them (Gosline, spine base remains to determine that the spine 1941). is not much thicker than the first soft-ray base. Like S. planiceps the gill membranes of the This is similar to the slender spine of S. plani- holotype are separate and non-overlapping to ceps, as are the overall form of the fin and its about the level of the rictus (Fig. 8B). We count dark pigmentation as originally illustrated (Fig. 14 branchiostegal rays on the right side of the 7). Eigenmann and Pearson (1924) stated that holotype. We have found relatively high bran- the dorsal fin of the holotype originates in the chiostegal counts of 13-15 in Sorubimichthys, third fifth of the body. This is a simple error as Pseudoplatystoma, Sorubim, some Brachyplatysto- shown by their figure and our reexamination; ma, and Goslinia, whereas most pimelodids have the dorsal fin originates in the second fifth of fewer branchiostegal rays. the standard length as in S. planiceps (predorsal Contrary to the statement of Eigenmann and length ca 38% of SL). Pearson (1924) the anterior nostril is not at the Eigenmann and Pearson (1924) reported the margin of the snout but a little behind it (Fig. anal count of P. manni as uncertainly 13; we 8A). This minor error is likely the result of the count 14 anal-fin rays including all rudiments. downward distortion of the snout noted earlier. In our sample of S. planiceps juveniles (greater Eigenmann and Pearson noted that the jaw teeth than about 14 mm) and adults the anal-fin rays of Pteroglanis manni are sharp, conical and in vary from 14-16 (modally 16). bands. In addition we find a few scattered teeth In the holotype of P. manni, as in S. planiceps, on the palate. We also find this early state of the eye is centered before the middle of the palatal dentition development in 15-21 mm LUNDBERG ET AL.-REVIEW OF SORUBIMICHTHYS 341

2.3- 22 L 21- A *1'* ..9 B 02.0 0 0 G 1.9 17 0 p t1- L 1t R t7 0 is E te- 0~~~~~ G t4 0 15 1.3- H Ro 141 EUE t2' 0 A t- D to- ? NA t L o * N 0.7 * N 07 G 0.6 0.6 T o05 La.s H -S 0.4. 0p 0.3' G 03 0.2 02 T o.1 H Q01 U0U I I I I I I I I I I I I I 0.o, , , , , , , O t1.1 13 1.4 5 1.6 7 1t to1. 2.0 21 22 23 2.4 25 2.6 27 0.0 0.2 0.4 0.6 0. 10 t2 14 t6 t' 2.0 22 2.4 LOG STANDARDLENGTH LOG PREDORSALFIN LENGTH

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Fig. 9. A-D) Bivariate scatterplots of log transformed measurements of Sorubimichthysplaniceps (closed circles) and Pteroglanis manni (star); E) Principal Components Analysis ordination of Sorubimichthysplaniceps (closed circles) and Pteroglanis manni (star). 342 COPEIA, 1989, NO. 2 specimens of S. planiceps (Fig. 5A). As noted caudal peduncle depth and length (PEDUNC- earlier the distinctive form and extent of the DEP, PEDUNCLEN). Measurements taken on band of palatal teeth in S. planiceps (Figs. 2, 5B) the smaller S. planiceps (<21 mm) included the is not developed until the fish are between 113 first seven above. Length of the projection of and 200 mm. the upperjaw beyond the mandibular symphysis In the original description the pelvic fin was (JAWLEN) was taken on all fish of 39 mm and stated to be about half the head length. This less. Bivariate scatterplots of representative log cannot be confirmed because of fin-ray break- transformed measurements for all fish show the age, but the pelvics in our comparably sized general similarity of the holotype to S. planiceps specimens of S. planiceps are approx. 50% of the (Fig. 9). Because of the very large difference in head length. The very dark pigmentation of the size, and the one order of magnitude gap be- pelvics of the holotype is preserved as another tween the very small size and large size groups point of similarity with juveniles of S. planiceps. of S. planiceps, quantitative comparisons of the An important feature of the holotype that was holotype were made separately. For specimens well illustrated (Fig. 7) and noted by Eigenmann of S. planiceps exceeding 21 mm, linear regres- and Pearson (1924) is the anterior membranous sions and 95% confidence intervals were com- continuation of the adipose fin. They did not puted for the following paired measurements: comment on similar but lower fin fold mem- PD:SL, HEAD1:PD, HEAD2:PD, HEADW:PD, branes of the caudal-fin lobes that almost reach SNOUT:PD, INTEROC:PD, PELVIC-CAUD: the adipose and the anal fin bases. These mem- PD, EYE-PECT:PD, AFB:PD, BODYDEP:PD, branes are thin and mostly transparent or hya- EYE:SNOUT, PEDUNCDEP:PEDUNCLEN. line with a few scattered, superficial melano- For specimens less than 39 mm, linear regres- phores. An essentially identical set of fin fold sions and 95% confidence intervals were com- membranes is present in the 21 mm juvenile S. puted for the following paired measurements: planiceps. The fin fold membranes further sug- PD:SL, HEAD1:PD, HEADW:PD, SNOUT: gest that the holotype is a juvenile fish. The fin PD, INTEROC:PD, JAWLEN:SNOUT. The folds are reduced by 33 mm. measurements of the holotype were within the Today, the overall coloration of the holotype 95% confidence band for 11 of the 18 contrasts, appears to be considerably darker than shown and for the remaining seven contrasts the ho- in the original illustration. The pattern of me- lotype was just outside the confidence intervals, lanophores remains, however, and matches and in no case was the holotype outside by more closely that of the 21 mm specimen of S. plan- than some individuals of S. planiceps. These re- iceps described earlier. sults are consistent with the other data indicat- Morphometric comparisons between the ho- ing the overall similarity of the holotype to S. lotype and S. planiceps were made to further planiceps. assess their overall similarity. Two series of S. Also, a Principal Components Analysis was planiceps were used in these comparisons: 14 performed on the covariance matrix of six log small juveniles ranging from 10-39 mm, and transformed measurements on the juveniles and 10 larger subadult/adult fish ranging from 209- the holotype: SL, PD, HEAD1, HEADW, 475 mm. Fourteen measurements were taken SNOUT, INTEROC. The first component was on the holotype, the 21, 33, 39 mm, and the 10 a "body size" factor with all variables loading larger S. planiceps: standard length (SL), pre- positively and about equally. An ordination (Fig. dorsal length (PD), head length measured to the 9E) was made by plotting the second and third end of the operculum membrane (HEAD1), component scores. The second component con- head length measured to the end of the bony trasts predorsal length to interorbital width and opercle (HEAD2), head width measured across snout length, whereas the third contrasts head the bases of the cleithra just behind the gill width and PD. The holotype of P. manni lies membrane (HEADW), snout length (SNOUT), near the centroid in this multivariate ordina- interocular distance (i.e., between the dorsal tion, indicating that on the basis of the covaria- margins of the orbital rims) (INTEROC), dis- tion of these body dimensions P. manni is not tance between the pelvic fin origin and caudal at all eccentric relative to juveniles of S. plani- fin base (PELVIC-CAUD), least distance be- ceps. tween the eye and pectoral origin (EYE-PECT), An additional piece of information involves anal fin base length (AFB), body depth at dorsal the behavior and habitat of the holotype which fin origin (BODYDEP), eye diameter (EYE), according to Mann's field notes was collected LUNDBERG ET AL.-REVIEW OF SORUBIMICHTHYS 343 as it hovered in a bed a vegetation. These notes central interest to us. R. M. Bailey called our match our field observations of juvenile S. plan- attention to Fowler's Pteroglanis. Critical input iceps. from J. N. Baskin and D. E. Stewart improved In summary, we have observed many detailed this study greatly, but we remain solely respon- similarities between the holotype of P. manni sible for any errors of fact and interpretation. and juveniles of S. planiceps and we have found J. R. Bailey, A. H. Bornbusch and L. A. McDade no differences that cannot be explained as pres- gave us several helpful suggestions. ervation artifacts or expected intraspecific vari- Support for JGL to travel to Venezuela in ation. We conclude that these nominal taxa are connection with this study came from the United conspecific and P. manni should be treated as a States Information Service Council for the In- subjective junior synonym of S. planiceps. ternational Exchange of Scholars through an We want to note the significance of the re- American Republics Research Grant and from cently discovered diversity of early life history the Duke University Research Council. Support stages of pimelodid catfishes (Mago-Leccia et for FML and PN to collect and study juvenile al., 1986). Based on the small amount of pub- pimelodid catfishes was provided by the Vene- lished descriptive work, the picture of siluri- zuelan Consejo Nacional de Investigaciones form early development is one of relatively "di- Cientificas y Tecnologicas (CONICIT), Pro- rect" development to the adult form (references yecto SI-1500. in Fuiman, 1984). Pimelodid catfishes exhibit some exciting exceptions to this pattern that LITERATURE CITED will surely play an important role in understand- their and ing systematics ecology. BLEEKER,P. 1862. Atlas ichthyologique des Indes Orientales Nberlandaises.Vol. 2. Frederic Miller, Amsterdam,The Netherlands. COMPARATIVE MATERIAL OF EIGENMANN,C. H., ANDR. S. EIGENMANN.1888. Pre- PIMELODIDAE JUVENILE liminarynotes on South American Nematognathi. Institutional abbreviations are as listed in Leviton et al., 1985. The Proc. Calif. Acad. Sci., Ser. 2, 1:119-172. following material is deposited in the collection of fishes at the , AND . 1890. A revision of the South MBUCV: Brachyplatvstomajuruense MBUCV-J-161-163; B. rousseauxi American Nematognathi or cat-fishes. Occ. Pap. MBUCV-J-14,29-131; Hemisorubin platyrhynchos MBUCV-J-87,88; Calif. Acad. Sci. 1:1-509. Megalonema platycephalum MBUCV-J-149; Megalonema sp. MBUCV-J- 151; Phractocephalus hemiliopterus MBUCV-J-37,84; blochi , AND N. E. PEARSON.1924. Pteroglanis, Ptero- MBUCV-J-46,59,62,69,70,74,76,115-117,119; P. pictus MBUCV-J- glanis manni, p. 9-10. In: The fishes of the eastern 73,111,112,153,156-158; Pinirampus pirinampu MBUCV-J-105- of the Andes. I. The fishes of the Rio Beni cf. Pseu- slope 110,124,125; Pseudopimelodus apurensis MBUCV-J-56,57,81; Basin, collected the Mulford doplatystomafasciatum MBUCV-J-8,41-45,90; P. tigrinum MBUCV-J-89; Bolivia, by Expedition. Sorubim lima MBUCV-J- 1,3,38,39,52,60,63,68,91,102,103; Sorubimich- N. E. Pearson (ed.). Ind. Univ. Stud. 11(64):1-83. thys planiceps MBUCV-J-78,85,86,101,113,128. FOWLER,H. W. 1934a. Zoological results of the third Additional uncataloged Venezuelan or Colombian material of juve- de SchauenseeSiamese expedition, Part I.-Fishes. nile catfishes at DU: Brachyplatystomaspp., Megalonemacf. platycephalum, Proc. Acad. Nat. Sci. Phila. 86:67-163. Megalonemasp., Pimelodusblochi, P. pictus, Pinirampus pirinampu, Sorubim; and one juvenile of Sorubimichthysplaniceps from Peru deposited at the . 1934b. Zoological results of the third de FMNH (FMNH96147). Schauensee Siamese expedition, Part V.-Addi- tional fishes. Ibid. 86:335-352. . 1951. Os peixes de aqua doce do Brasil.Arq. ACKNOWLEDGMENTS Zool. Estado S5o Paulo 9:1-400. FUIMAN,L. 1984. and re- We thank the staff of the of Ich- Ostariophysi:development Department lationships,p. 126-139. In: Ontogeny and system- thyology and the Library of the California atics of fishes-an international symposiumdedi- Academy of Sciences for permission to study cated to the memory of Elbert Halvor Ahlstrom. the type of Pteroglanis manni, and their copy of H. G. Moser, W. J. Richards,D. M. Cohen, M. P. Spix and Agassiz (1829-1831). The staff of the Fahay, A. W. Kendall, Jr. and S. L. Richardson Philadelphia Academy of Natural Sciences made (eds.). 15-18 Aug. 1983, La Jolla, California. American of and their copy of Spix and Agassiz available for pho- Society Ichthyologists Herpetol- The staffof the Duke Rare ogists (ASIH), Gainesville,Florida. tography. University T. 1870. Fishes from the Maranonand Book Room allowed us to their of GILL, Napo study copy Rivers. Proc. Acad. Nat. Sci. Phila. 1870:92-96. Seba's Planches. We are indebted to especially GOSLINE,W. A. 1941. Synopsis of the genera of pi- G. Howes, T. Mo, M. Kottelat, andJ.-P. Haenni melodid catfishes without a free orbital rim. Stan. for their generosity in providing information Ichthy. Bull. 2(3):83-88. on and photographs of specimens that were of . 1945. Catalogo dos nematognatos da agua- 344 COPEIA, 1989, NO. 2

doce da America do Sul e Central. Bolet. Museu MIRANDORIBEIRO, A. 1911. Fauna Brasiliense. Pei- Nac., Nov. Ser., Rio de Janeiro 33:1-138. xes. V(A). Eleutherobranchquios Aspirophoros. GUNTHER,A. 1864. Catalogue of the fishes in the Arq. Mus. Nac. Rio de Janeiro 16:1-504. collection of the British Museum. Vol. 5. Trustees SCHULTZ,L. P. 1944. The catfishes of Venezuela of the British Museum, London, England. with descriptions of thirty-eight new forms. Proc. . 1872. Notice of a large siluroid (Platystoma U.S. Nat. Mus. 94(3172):173-388. gigas) from the upper Amazons. Ann. Mag. Nat. SPIX,J. B. VON, ANDL. AGASSIZ.1829-1831. Selecta Hist., Ser. 4, 10:449-450. genera et species piscium quos in itinere per Bra- LEVITON,A. E., R. H. GIBBS, JR., E. HEALAND C. E. siliam, annis MDCCCXVII-MDCCCXX jussu et DAWSON. 1985. Standards in herpetology and ich- auspiciis Maximiliani Josephi I. Bavariae regis au- thyology: Part I. Standard symbolic codes for in- gustissimi peracto colleget et pingendos curavit Dr. stitutional resource collections in herpetology and J. B. de Spix, digessit, descripit et observationibus ichthyology. Copeia 1985 (3):802-832. anatomicus illustravit Dr. L. Agassiz, praefatus est LEVRAULT,F. G. 1827-1828. Planches de Seba. Vol. et edidit itineris socius Dr. F. C. Ph. de Martius. 3. Paris, France. Munich, Germany. LUNDBERG, J. G., O. LINARES, P. NASS AND M. E. STEWART, D. J. 1986. Revision of Pimelodina and ANTONIO. 1988. Phractocephalus hemiliopterus(Pi- description of a new genus and species from the melodidae, Siluriformes) from the upper Miocene Peruvian Amazon (Pisces: Pimelodidae). Copeia Urumaco Formation, Venezuela: a further case of 1986:653-671. evolutionary stasis and local extinction among South WHITEHEAD, P. J. P., AND G. S. MYERS. 1971. Prob- American fishes. J. Vert. Paleo. 8(2):131-138. lems in nomenclature and dating of Spix and Agas- , AND L. A. MCDADE. 1986. On the South siz's Brazilianfishes (1829-1831).J. Biblio. Nat. His- American catfish Brachyrhamdiaimitator Myers (Si- to. 5(6):478-497. luriformes, Pimelodidae), with phylogenetic evi- dence for a large intrafamilial lineage. Acad. Nat. (JGL) DEPARTMENT OF ZOOLOGY, DUKE Sci. Phila., Not. Nat. 463:1-24. UNIVERSITY, DURHAM, NORTH CAROLINA P. NASS AND O. CASTILLO.1986. MAGO-LECCIA,F., 27706 AND (PN, FML) INSTITUTO DE adultos de de la familia Larvas, juvenilies y bagres ZOOLOGIA TROPICAL, UNIVERSIDAD CENTRAL Pimelodidae Siluriformes) de Venezue- (Teleostei, DE VENEZUELA, CARACAS, VENEZUELA. Ac- la. Informe Final SI-1500, CONOCIT, Proyecto 5 1988. Caracas, Venezuela. cepted Aug.

Copeia, 1989(2), pp. 344-348

Spatial and Temporal Variation of Mitochondrial DNA Haplotype Frequencies in the Striped Bass (Moronesaxatilis) 1982 Year Class

ROBERT W. CHAPMAN

This report summarizes mitochondrial DNA variation in 195 specimens of striped bass, Morone saxatilis, from the 1982 year class. Specimens were collected from 1984-87 from three spawning areas and from wintering grounds within Chesapeake Bay. The analysis suggests that striped bass males have little fidelity to natal spawning grounds, while females appear to have a stronger homing instinct.

T HE striped bass, Morone saxatilis, has sup- the commercial harvest. Berggren and Leiber- ported important commercial and rec- man (1978) estimated that 70+ % of the coastal reation fisheries along the Atlantic and Gulf of (oceanic) harvest of striped bass was derived from Mexico coast of the US. Historically, Chesa- Chesapeake stocks. Studies of migratory pat- peake Bay has been a major nursery and fishing terns of M. saxatilis in Chesapeake Bay suggest ground for this species contributing 50+% of that following birth in tidal freshwaters, young

? 1989 by the American Society of Ichthyologists and Herpetologists