Mimagoniates microlepis (Steindachner, 1877)

Menezes ¹, Naércio A. & Weitzman ², Stanley H., 2009, Systematics of the Neotropical fish subfamily Glandulocaudinae (Teleostei: Characiformes: Characidae), Neotropical Ichthyology 7 (3), pp. 295-370 : 355-364

Mimagoniates microlepis (Steindachner, 1877) Figs. 77-79

Paragoniates microlepis Steindachner, 1877: 33 (type locality: “Bäche in der Nähe von Rio de Janeiro, rio dos Macacos”; although this volume is for the year 1876 and often so cited the date of publication was 1877). - Eigenmann & Eigenmann, 1891: 57 (listed).- Eigenmann, 1910: 441 (listed).

Coalurichthys iporangae Miranda-Ribeiro, 1908 (unpaginated); type locality: “Ribeirão das Pedras, Iporanga”; spelling error for generic name; see generic synonymy for Mimagoniates .

Coelurichthys iporangae Eigenmann, 1910: 428 (listed). Eigenmann 1914: 42 (listed). - Nichols, 1913: 152 (in key to species). - Travassos, 1951: 42 (listed). - P. Miranda Ribeiro, 1955: 390 (listed).

Coelurichthys microlepis Rachow, 1927: 17 (aquarium description). Rachow in Holly Meinken & Rachow, 1950a: 755 (in synonymy). - Nelson, 1964a: 62, 63, 68 (anatomy; systematics; courtship behavior). - Géry, 1966: 228, 230 (discussion and in key). - Géry, 1977: 362 (listed in discussion). - Sterba, 1987: 69 (aquarium description).

Mimagoniates microlepis Rachow, 1928: 16 (aquarium description). - Myers, 1928: 120 (listed). - Eigenmann & Myers, 1929: 491 (redescription; figure). - Innes, 1935: 123 (aquarium description). - Rachow, in Holly, Meinken & Rachow, 1950a: 755 (aquarium description; synonymy of aquarium literature). - Schultz, 1959: 10 (in part only; listed M. tenuis as synonym; of other specimens examined and assigned to M. microlepis, USNM 86296 is M. barberi, USNM 177703 are M. lateralis ). - Weitzman & Fink, 1985:98, 106 (discussion of caudal pump structure; listed in specimens examined).-Weitzman et al., 1988:395, 396,409,412 (anatomy of caudal gland; discussion of phylogeny and biogeography; figure). - Menezes & Weitzman, 1990: 381, 386, 388, 389 (phylogeny; in key; anatomy of caudal gland). - Guimarães et al., 1995: 185-189 (cytogenetic studies). - Weitzman et al., 1996a: 203-204 (breeding; ecological data; Weitzman et al., 1996b: 195-196, 199-209 (photograph, illustration of tail fin; distribution, ecological data, relationships; courtship behavior). - Malabarba & Weitzman, 1999: 84 (listed in discussion). - Malabarba & Weitzman, 2000: 279 (listed in discussion). - López et al., 2002: 59 (listed in type catalog). - Weitzman, 2003: 226 (maximum length; distribution; remarks and references). - Ingenito et al., 2004: 26, 27, 29 (in key; comments on collecting activities and origin of the species in rio Iguaçu) - Sant’Anna et al., 2006: 42-45 (first record in rio Tibagi basin; comments on distribution). - Oyakawa et al., 2006: 152 (photograph; diagnosis; size, distribution; ecological data). - Langeani e t al., 2007: 184 (listed in upper rio Paraná). - Torres et al., 2007: 1-7 (chromosomes; molecular data; evolutionary trends). - Menezes et al., 2007: 125 (photograph; distribution; systematic status; synonyms; ecology; conservation). - Menezes, 2007: 39 (listed in catalog; distribution). - Menezes et al., 2008: 37-40, 43-45 (photograph; distribution; molecular analysis; discussion of relationships and biogeography).

Coelurichthys lateralis Myers in Eigenmann & Myers, 1929: 491, 492 (type examined and erroneously referred it to M. microlepis ). - Rachow in Holly, Meinken & Rachow, 1950a: 755 (referred species to M. microlepis following Myers, in Eigenmann & Myers, 1929: 491, 492. - Schultz, 1959: 11 (followed Myers, in Eigenmann and Myers, 1929: 491, 492). - Nelson, 1964a: 65 (in part; found female type of C. lateralis difficult to identify and could not decide whether it was C. microlepis or C. tenuis but if latter, C. lateralis would have priority because of page precedence).

Coelurichthys tenuis Myers , in Eigenmann & Myers, 1929: 491, 492 (type examined and erroneously referred it to M. microlepis ). - Rachow, in Holly Meinken & Rachow, 1950a: 755 (followed Myers, in Eigenmann & Myers 1929: 491, 492 in referring this nominal species to M. microlepis ).

Mimagoniates inequalis Schultz, 1959: 63 (in part only, one specimen of M. microlepis, USNM 177704, was misidentified as M. inequalis ).

Mimagoniates cf. microlepis Géry, 1964: 6 , (discussion; osteology of caudal and anal fins).

Mimagoniates sp. C , Weitzman et al., 1988: 411-412 (osteology of caudal fin).

Diagnosis. Mimagoniates microlepis can be distinguished from all the other species of Mimagoniates except M. rheocharis by the presence of a fully developed caudal-fin ray pump and hooks on some principal caudal-fin rays. In addition to having caudal-fin hooks spiny (vs. sturdy in M. rheocharis ), M. microlepis differs from M. rheocharis by the number of horizontal scale rows between dorsal- and anal-fin origins (15-18 vs. 17-22 for M. rheocharis ), scales around caudal peduncle (15-18 vs. 19-23 for M. rheocharis ), and coloration as discussed in the diagnosis of M. rheocharis .

Description. Table 12 presents morphometrics of the lectotype, paralectotype and topotypes. Except where noted, the entire description refers to the population sample represented by specimens from rio Macacu, the type locality or adjacent tributaries flowing into this river. These collections were treated statistically as one population sample in an attempt to represent the species. Variations of meristic and morphometric data within the range of M. microlepis are discussed where appropriate.

Body compressed, relatively deep, especially near dorsal-fin origin; body deepest at vertical through anal-fin origin. Predorsal body profile relatively arched in adult males, less so in adult females and immatures which have predorsal profile gently convex. Body profile elevated at dorsal-fin origin, less so in females and juveniles. Dorsal profile of body nearly straight along dorsal-fin base to adipose fin. Body profile posterior to adipose fin somewhat concave dorsal to caudal

Males Females and juveniles

Characters

Holotype N Range Mean SD N Range Mean SD dif.

Standard length 47.5 28 30.0-47.5 31.4 15 19.0-30.4 24.0

Depth at dorsal-fin origin 30.9 28 24.5-30.9 26.5 1.6 15 21.2-26.6 24.4 1.7 + Snout to dorsal-fin origin 60.2 28 57.5-65.0 61.6 1.3 15 59.4-63.8 61.3 1.3 - Snout to pectoral-fin origin 26.5 28 23.3-26.7 25.3 1.0 15 23.4-26.4 24.8 1.1 - Snout to pelvic-fin origin 43.4 28 42.4-45.6 44.3 0.8 15 40.1-44.7 43.3 1.5 - Snout to anal-fin origin 56.8 28 53.0-58.6 56.5 1.3 15 54.2-58.4 56.1 1.4 - Caudal peduncle depth 14.7 28 11.4-14.7 13.2 0.9 15 09.4-13.9 11.2 1.2 + Caudal peduncle length 9.3 28 06.2-09.2 07.8 0.8 15 07.8-09.1 08.4 0.4 - Pectoral-fin length 21.7 28 20.4-25.8 23.2 1.4 15 20.0-24.5 21.8 1.3 - Pelvic-fin length 13.9 28 12.6-16.9 14.8 1.1 15 10.5-14.6 12.6 1.2 + Dorsal-fin base length 14.1 28 12.0-15.9 14.2 0.9 15 10.5-14.2 12.4 1.0 - Dorsal-fin height 38.3 27 25.2-38.3 31.3 3.2 15 19.1-24.4 21.8 1.2 + Anal-fin base length 36.4 28 35.7-40.0 37.2 1.2 15 35.1-39.4 36.9 1.3 - Anal-fin lobe length 30.1 26 16.0-20.4 18.4 1.5 15 17.7-21.0 19.1 0.9 -

Eye to dorsal-fin origin 51.2 28 45.8-51.2 49.0 1.3 15 46.1-50.3 47.8 1.0 - Dorsal-fin origin to caudal-fin base 42.9 28 37.4-44.5 41.2 1.8 15 38.0-42.7 40.2 1.3 - Bony head length 23.4 28 22.2-26.3 24.5 1.0 15 22.8-25.4 24.0 0.8 - Horizontal eye diameter 34.2 28 34.2-41.6 37.4 2.3 15 34.0-41.6 38.0 2.7 - Snout length 24.3 27 21.4-25.2 23.6 1.1 15 21.2-25.0 22.7 1.1 - Least interorbital width 34.2 28 32.0-36.6 34.3 1.4 15 32.7-36.9 35.2 1.3 - Upper jaw length 45.9 27 41.9-46.8 44.6 1.2 15 41.5-46.8 44.1 1.5 -

peduncle. Dorsal-fin origin nearer to caudal-fin base than to snout tip. Ventral profile of body strongly convex in adult males from tip of lower jaw to origin of pelvic fin, less strongly convex in females and juveniles. Abdominal profile in adult males slightly concave to anal-fin origin, straight or nearly so in females and juveniles. Body profile along anal-fin base in males slightly concave along base of anterior lobe of anal fin; straight along base of remainder of fin in males and along entire anal-fin base in females and juveniles. Ventral profile of caudal peduncle convex in adult males due to strong development of anterior 5 or 6 procurrent ventral caudal-fin rays included in that profile; slightly concave or nearly straight in females and juveniles.

Lower jaw protruding, slightly beyond upper jaw. Lower jaw of adult males thick and heavy compared to that of females and juveniles. Mouth angled posteroventrally. Maxilla long, extending to point ventral to horizontal through ventral border of eye. Maxilla extends posteriorly to a point anterior to vertical line drawn through anterior border of pupil.

Dorsal-fin rays ii, 8 in all specimens, n = 41; posterior ray not split to its base and counted as 1 ray. Anal-fin unbranched rays iv or v, usually iv, branched rays 26-31 (27), 28.2, n = 42; posterior ray split to its base and counted as 1 ray. Anal fin with moderately developed lobe anteriorly ( Figs. 77-79 and 81 ); lobe includes last unbranched ray and first 5 branched rays.Anal fin of sexually mature males with bilateral hooks, 1- 3, usually 1 on each side, on posterior most unbranched ray ( Fig. 81 ). Usually anterior 7-8 branched fin rays with bilateral hooks, 1 set for each ray. Pectoral-fin unbranched ray i in all specimens, branched rays 8-11 (9), 9.7, n = 42. Posterior tip of first pectoral fin extends posteriorly slightly beyond origin of pelvic fin. Pelvic fin rays 7 in all specimens. Pelvic fin with first ray branched in adult males and females ( Fig. 83 ), unbranched in juveniles of both sexes. Sexually mature, large adult males with over 200 hooks on each pelvic fin distributed as shown in Figs. 83 and 84 .

Principal caudal-fin ray count 10/ 9 in all specimens, n = 42. Fin rays modified in association with caudal pheromone pump as in Fig. 85 . Caudal-fin rays 8 to 12 with small bony hooks.

Scales cycloid, with few radii along posterior border. Terminal scale of modified caudal-fin series without exaggerated radii ( Fig. 85 ).

Lateral line incomplete, perforated scales = 6-9 (6), 7.1, n = 23. Lateral series scales 40-46 (43), 42.7, n = 21. Predorsal scales 21-24 (23), 22.9, n = 24. Scale rows between dorsal- and anal-fin origins 13-16 (14), 14.5, n = 34. Scale rows around caudal peduncle 15-17 (16), 16.3, n = 12.

Premaxillary teeth in 2 irregular almost undistinguishable rows and summary count for teeth on left premaxilla are given as a single count ( Fig. 86 ). Larger and smaller teeth tricuspid in all large specimens, sometimes in smaller specimens small teeth bicuspid or conical. Total count premaxillary teeth 7-11 (9), 8.8, n = 41. Maxillary teeth 3-7 (5), 4.09, larger specimens usually with more teeth counts, n = 41. All maxillary teeth tricuspid in large specimens; small specimens with posterior maxillary teeth often conical ( Fig. 86 ). Dentary with 4 large anterior tricuspid teeth, followed by smaller posterior teeth 4-11 (9), 7.5, almost always more teeth in largest specimens; anterior small dentary teeth of posterior series tricuspid, posterior ones conical, n = 41 ( Fig. 86 ). Maxillary and dentary teeth shaped much like premaxillary teeth.

Vertebrae 37-39 (38), 38.2, n = 42. Dorsal limb gill rakers 5- 7 (6), 6.2, n =41; ventral limb gill rakers 10-12 (12), 11.2, n = 41. Branchiostegal rays 4, in 11 cleared and stained specimens, 3 rays originating on anterior ceratohyal and 1 ray from posterior ceratohyal.

Color in alcohol. See Figs. 77-79 for preserved color pattern of males and females. Some variation occurs in color pattern of population samples from geographically isolated areas within species range. Following basic color pattern of specimens from the type locality characteristic of all specimens available. Body pale to medium brown, pale yellowish-brown ventrally, much darker dorsally. Humeral spot slightly vertically elongate, in both sexes, obscured by dark lateral stripe in some specimens. Dark lateral body stripe somewhat diffuse anteriorly, darker posteriorly, but uniformly dark in some specimens of both sexes. Stripe extending from humeral spot posteriorly onto dorsal region of ventral caudal-fin lobe and ventral portion of dorsal caudal-fin lobe. Lateral stripe extends over all caudal gland structures. Stripe especially dark on principal rays 10, 11 and 12, less so on ray 13. Remainder of caudal fin dusky due to presence of scattered dark chromatophores. Dorsal border of first principal caudal-fin ray and ventral border of nineteenth principal caudal-fin ray black. Middorsal body surface black, forming narrow stripe extending from supraoccipital region to base of dorsal procurrent rays of caudal fin. Indistinct narrow dark stripe between dorsal black stripe and lateral stripe posteriorly from dorsal-fin origin to caudal base. Remainder of dorsal body surface ventral to lateral body stripe pale brown.

Pectoral and pelvic fins dark, dorsal and anal dusky with scattered dark chromatophores along fin rays and membranes. Anal fin with dark elongate stripe running length of fin. Stripe width about one-third height of anal fin anteriorly and about one-fourth fin’s height posteriorly; bordering edge of fin from sixth or seventh branched rays to posterior tip of last rays. Stripe of nearly uniform width throughout its length. Stripe somewhat darker anteriorly in sexually mature males, especially dorsal to relatively hyaline anterior tip of anal-fin lobe. Dorsal-fin with horizontal dark stripe in adult males and females extending posteriorly from about mid-length of anterior elongate unbranched ray to posterior tips of two terminal dorsal-fin rays. Stripe usually narrow, less than one-eighth maximum height of dorsal fin. Width and intensity of stripe variable depending on sex and sexual maturity. Preserved males sometimes with anterior portion of stripe diffuse. Adipose fin dusky with scattered dark chromatophores.

Head light to dark brown around mouth and on dorsal surface of snout, between eyes, dorsal portion of cranium and nape. Tip of lower jaw light to dark brown. Portion of head posterior to infraorbitals and extending ventrally from parietal region across dorsal opercular region dark brown. Dark area continues ventrally across posterior region of opercle to just reach interopercle and looks similar in form as humeral spot. Iris dark brown, dorsal to pupil and silvery elsewhere. Infraorbitals silvery where guanine preserved, pale yellowish brown when guanine absent. Dark brown chromatophores scattered evenly through circumorbital area. Anterior area of opercle, all of preopercle, and branchiostegal rays silvery or pale brown, without much dark brown pigment.

Color in life. Life colors described here taken from a 35 mm color slide of an adult male collected in rio Sagrado system photographed just after capture from clear slightly turbid water stream, baía de Guaratuba, Paraná. Sides of body pale silvery blue. Broad lateral body stripe somewhat deeper silvery blue from humeral spot to caudal peduncle. Just dorsal to silvery blue color of body sides, dorsum with narrow dark brown line extending from parietal region of head to caudal peduncle, black with blue pigment on procurrent rays. Lateral portion of dorsum between narrow brown line and lateral body stripe light brown. Lower jaw and upper part of head above eye dark with blue pigment. Infraorbitals and ventral part of preopercle and opercle silvery white with scattered blue chromatophores. Dorsal part of preopercle and opercle dark with scattered blue and black chromatophores. Ventral portion of head from about vertical through anterior border of eye and abdominal region yellowish. Lower part of body ventral to lateral stripe silvery with scattered blue and yellow chromatophores. Basal part of anal fin yellow with scattered blue chromatophores. Caudal fin hyaline with blue pigmentation especially evident on middle caudal-fin rays. Ventral most unbranched principal caudal-fin ray distally black. Remainder of black pigment of caudal fin in preserved specimens. Anal fin with distal portion of fin rays posterior to anterior anal-fin lobe dark obscured by blue pigmentation, forming stripe along margin of fin. Distal region of anterior anal-fin lobe yellow, proximally bordered by black pigment mixed with yellow and blue. Approximately distal one-half of pelvic fin yellow; remaining proximal portion of fin hyaline. Distal one-half of pectoral fins yellow, proximal half hyaline. Dorsal fin hyaline distal and proximal to longitudinal black longitudinal stripe partly obscured by yellow and blue pigmentation.

Sexual dimorphism. Females lack a caudal pheromone pump organ and pelvic-fin and anal-fin hooks described above for males. Life colors of females more subdued than that of sexually mature males. Among the characters showing significant differences between males and females as indicated in Table 12, only dorsal-fin height proved to be statistically different through regression analysis. Figure 88 clearly shows a significant difference expressed by the slopes of males and females indicating a divergence at all sizes.

Distribution. Mimagoniates microlepis is widely distributed in the coastal area from southern Bahia to northern Rio Grande do Sul and also in the upper rio Iguaçu and rio Tibagi, upper rio Paraná basin, Brazil. See figure 3 in Menezes et al. (2008).

Ecology. Weitzman et al. (1988: 413) mentioned that M. microlepis occurs in clear running waters of small to large streams and is quite common and most abundant in forested areas near the shore. Subsequently Menezes & Weitzman (1990: 423) pointed out that it is rarely found in black acid waters. More recent collections of this species obtained from eastern and southeastern Brazilian coasts, however, indicated that it is more common in blackwaters than previously thought. It has been recently collected in small black water streams with substrate consisting of clay, rocks and sand as well as in clear water streams in Santa Catarina. In most other places both in eastern and southern Brazil, M. microlepis was caught in slow moving clear water streams and small ponds, even in areas where the original Mata Atlântica vegetation was removed. In the small streams the water was cool and rocks, sand, mud and twigs fallen from isolated trees were usually found on the bottom.

Data from Costa (1987) indicate that M. microlepis lives near the surface and feeds mostly on terrestrial arthropods. Sabino & Castro (1990) found that it is primarily insectivorous and that 73.6% of its diet consists of items that fall into the water, especially insects (63.15%) and arachnids (10.5%). Essentially the same results were obtained by Lampert et al. (2003) who concluded that M. microlepis is insectivorous regardless of size and sex.

Remarks. The discussion of meristic and morphometric differences as well as possible hybrid origins involving M. microlepis , M. rheocharis and M. inequalis in Menezes & Weitzman (1990: 416-421) are not repeated here. For further comments on relationships of M. microlepis see the “Phylogeny” section.

Anal-fin branched ray counts, number of scales (horizontal scale rows on body, horizontal scale rows around caudal peduncle, lateral-series scales and predorsal scales), and number of vertebrae of Mimagoniates microlepis showed significant differences among population samples of the species ( Figs. 89 and 90 ). For the purpose of comparison, the samples were grouped within the biogeographical coastal subregions defined by Menezes (1988: 300). Comparing only samples included in the North Coastal Subregion with the samples included in the more southern upland areas of Paraná the respective ranges and medians are significantly different, with very little overlap in some cases. However, the same values for the intermediate samples included in the South Coastal and in the two Central Coastal subregions bridge the gap. Thus there is a pattern of latitudinal variation of characters than sharp differences that would justify the recognition of more than one species. The recent discovery of specimens identified as M. microlepis from the rio Tibagi basin ( Sant’Anna et al., 2006) flowing through the upper Paraná basin demonstrates that it is more widespread than previously indicated. The distribution of some isolated populations of M. microlepis is discussed by Menezes et al. (2008). Until a more detailed analysis of character variation within the range of M. microlepis can be performed, we prefer for the moment to consider it a widespread species represented by isolated populations.

No date of collection of the lectotype of M. microlepis, NMW 56534, was provided, but NMW catalog cites the collector as Steindachner and the date of receipt of specimens as 1874. Likely the specimens originated near town of Macacu (also called Cachoeiras de Macacu , approximately 22°26’S 42°49’W) in 1874 by Steindachner during Hassler Expedition , 1871-1872. Our observations indicate that species frequently occurs in small tributaries of rio Macacu and along shores of main river in emergent vegetation .

The holotype of Coelurichthys iporangae, USNM 92955 is apparently identical to that in photograph published by Miranda Ribeiro (1908: unpaginated). Both the specimen and photograph display the same bent body shape, the same wound to the upper portion of the abdominal cavity on the left body side. Miranda-Ribeiro’s original description is in the singular as though he was describing one specimen. However the number of specimens he had at hand is in some question. In the introduction to his study Miranda Ribeiro (1908) states “Both the Museu de São Paulo and the Museu Nacional have the material evidences of these studies”. We found a second specimen, MZUSP 408, 42.7 mm SL, collected by Krone from the ribeirão das Pedras, Iporanga, a tributary of the rio Ribeira de Iguape, São Paulo. No date of collection is given for this fish but we strongly suspect this specimen may have been collected with the type and probably sent to the Museu Paulista by Ricardo Krone, and subsequently transferred to the Departamento de Zoologia (now the Museu de Zoologia), Universidade de São Paulo. Miranda Ribeiro perhaps saw the specimen now at MZUSP but used only the specimen sent to the Museu Nacional, Rio de Janeiro in his original description. There is no absolute evidence that A. Miranda Ribeiro ever saw the MZUSP specimen. In any case this specimen appears not to have been previously discussed in the ichthyological systematic literature.

The fish here considered as holotype of C. iporangae, USNM 92955, cataloged on 11 Aug. 1933 as part of a collection donated to the USNM by George S. Myers. The fish had been obtained from the Museu Nacional, Rio de Janeiro by Eigenmann for his studies. In the USNM accession papers, (accession 124667), Myers listed this fish as a paratype and noted that he had earlier received the specimen from Eigenmann on exchange. Myers erroneously listed the collector as Alipio de Miranda Ribeiro rather than Ricardo Krone. This specimen was in turn listed by Eigenmann & Myers (1929: 491) as “Mus. Nac. Rio de Janeiro, type, number of specimens 1, size 51 mm, locality Iporanga, collector Haseman”. The references to Haseman and Miranda Ribeiro as collectors are clearly errors according to Myers in conversation with one of us (SW) in 1980.At the time of the publication of Eigenmann & Myers (1929) and the cataloging of this specimen at USNM, Myers had not seen Miranda Ribeiro (1908). Furthermore Myers in 1980 agreed with us that USNM 92955 is the holotype of C. iporangae . Based on the accession papers by Myers there is a notation in the USNM catalog that this specimen is a paratype. Myers had USNM 92955 specimen when he revised Eigenmann’s manuscript for publication in 1929 and kept the fish in his personal fish collections until he donated it to the USNM in 1933. The length of the specimen cited in Eigenmann & Myers (1929), 51 mm, is total length and agrees with the length of the holotype. This specimen was apparently not examined by L. P. Schultz.

Material examined. Lectotype. NMW 56534 : 1, male, 47.5 mm SL, Brazil, Rio de Janeiro State, “Bäche in der Nähe von Rio de Janeiro, rio dos Macacos” ( rio Macacu , a tributary to Baía de Guanabara on northeastern shore) . Paralectotype. NMW 56534 : 1, female, 41.0 mm SL, same data as lectotype . Topotypes from rio Macacu , Rio de Janeiro . MZUSP 14667 , 13 , young to juveniles, 19.0-30.0 mm SL, rio Guapiaçu, tributary of rio Macacu, near Cachoeiras de Macacu , R. M. C. Castro, 14 May 1977 ; MZUSP 14668 , 7 , males, SL 35.0- 40.7 mm SL ; MZUSP 14669 , 13 , young to small adults 12.5-37.7 mm SL ; MZUSP 26895 , 41 , young to juveniles, 13.1-26.9 mm SL, small tributary of rio Macacu about 1-2 km from town of Cachoeiras de Macacu, S. Weitzman, C. A. Gonçalves de Cruz and O. L. Peixoto, 27 Nov. 1979 ; USNM 249896 , 17 , young to small adults, 22.0- 35.4 mm SL ; USNM 254255 , 40 , young-small adults, SL 12.0- 30.8 mm . Holotype of Coelurichthys iporangae . USNM 92955 , adult male with well-developed caudal organ, 37.2 mm SL, Brazil, São Paulo, “Ribeirão das Pedras, Iporanga”, rio Ribeira de Iguape system, approximately 24°36’S 48°34’W . Topotypes. FMNH 54886 , 7 , young to small adult, 17.4 to 36.4 mm SL, Brazil, São Paulo, Iporanga, rio Ribeira de Iguape. Non-types. All collected in Brazil. Bahia State, município de Prado . USNM 279875 , 6 , young to immatures, 18.8-25.0 mm SL, first stream of rio Caí on dirt road between Cumuruxatiba and Itamaraju, stream locally called rio do Sul, 17°00’S 39°12’W ; MZUSP 53278 , 5 , young to immatures, 17.1- 23.2 mm SL ; USNM 279874 , 1 , immature, 16.5 mm SL, 17°01’S 39°12’W ; MZUSP 53279 , 1 , immature, 23.9 mm SL ; MZUSP 28814 , 3 , immatures, 15.4-22.2 mm SL, fourth stream flowing east at 26 km on dirt road north from Prado, rio Japurá , 17°10’S 39°14’W ; MZUSP 28813 , 26 , young to small adults, 18.1-29.8 mm SL, stream 18 km north from Prado on road to Cumuruxatiba , 17°10’S 39°14”W; USNM 276549 , 24 , young to small adults, 17.4-33.1 mm SL ; USNM 276550 , 2 , young adult female and male, 24.2-28.6 mm SL respectively ; USNM 276548 , 11 , immatures to small adults, SL 19.3-27.7 mm SL . MUSP 28812 , 11 , immatures to small adults, small stream about 5 km north of Prado, on road to Cumuruxatiba , 17°18’S 39°14’W ; USNM 249895 , 7 , immatures to small adults, 24.9-32.0 mm SL, rio Jacurucu at Prado, 17°21’S 39°13’W ; USNM 313657 , 1 , c&s, mature male, SL 35.4 mm. Following lots collected in Espírito Santo State . USNM 254257 , 20 immatures, 21.7-31.6 mm SL, rio Itaúnas at Itaúnas, 18°26’S 39°40’W ; MZUSP 26898 , 17 , immatures to small adults, 20.5-33.0 mm SL ; MNRJ 5675 , ribeirão do Engano, vale do Itaúnas, Instituto Oswaldo Cruz ; MNRJ 5679 , 82 , immatures to adults, 18.2- 41.3 mm SL ; MNRJ 10853 , 1 , immature, SL 24.6 mm, rio Barra Seca , between Linhares and São Mateus, approximately 19°02’S 40°05’W ; MNRJ 10855 , 22 , young to adults, 19.5-36.5 mm SL, município de Linhares , córrego Chumbado on road between Linhares and São Mateus ; MZUSP 26894 , 13 , immatures to adults, 25.3-42.2 mm SL município de Linhares, Reserva Florestal da Companhia do Vale do Rio Doce ; USNM 249885 , 11 , immatures to adults, 23.2-41.2 mm SL ; USNM 313656 , 2 , c&s, male, 41.2 mm SL, female, 26.4 mm SL ; USNM 249891 , 25 , immatures to adults, 22.4-24.8 mm SL MZUSP 26893 , 2 , adults, female, 28.4 mm SL and male 28.5 mm SL ; USNM 313655 , 2 , c&s, immature female, adult male, 27.3-31.2 mm SL respectively ; USNM 279873 , immatures 7, 22.4-31.7 mm SL, blackwater creek in Reserva Florestal da Companhia do Vale do Rio Doce ; MNRJ 10854 , 20 , immatures to small adults, 22.5-34.7 mm SL, Serra, town north of Vitória , 20°08’S, 40°18’W ; FMNH 15004 , 15005 , 15006 , 3 , maturing to adult males, 31.5-36.6 mm SL, Muniz Freire, sandy rocky clearwater stream one mile below town, a tributary to rio Itapemirim , approximately 20°28’S 41°24’W ; FMNH 54885 , 6 , maturing to adult males, 30.5-39.0 mm SL. Following lots collected in Rio de Janeiro State. See also lectotype, paralectotype and topotypes above . USNM 249884 , 4 immatures, 26.6-32.0 mm SL, small stream tributary to rio Macaé , adjacent to ruins of fazenda Torreão, 22°16’S 42°03’W ; MZUSP 26905 , 4 , small adults, 28.9- 30.5 mm SL ; USNM 249878 , 51 , immatures to fully mature adult females, males, 18.5-25.9 mm SL, município de Saquarema, small stream, rio Jundiá tributary to lagoa de Saquarema, on road Amaral Peixoto between Sampaio Correia and Bacaxá, 22°51’S 42°34’W ; MZUSP 26903 , 51 , young to mature adults, 16.8-22.6 mm SL ; MNRJ 26619 , 51 , young to mature adults, 18.5-25.4 mm SL ; USNM 249898 , 6 immatures, 22.2-26.4 mm SL, município de Saquarema, rio Mato Grosso into lagoa Saquarema, 5 km east of Sampaio Correa , 22°51’S 42°35’W ; USNM 257199 , 1 , adult female, 36.1 mm SL, aquarium fish grown up in aquarium ; USNM 249877 , 50 immatures, 13.7-33.2 mm SL, município de Saquarema, rio Seco , tributary to lagoa de Saquarema, along roadAmaral Peixoto between Sampaio Correia and Bacaxá, 22°51’S 42°32’W , MZUSP 26907 , 50 immatures, young to mature adults, 16.3-28.5 mm SL ; USNM 249875 , 2 , adult female, adult male, 29.0- 39.3 mm SL respectively, município de Saquarema, rio do Mato Grosso, tributary to rio Roncador which flows to lagoa Saquarema, 22°53’S 42°44’W ; MZUSP 26904 , 3 , adults, 30.0- 38.5 mm SL ; USNM 249900 , 33 , young to adults, 17.9-34.4 mm SL, município de Saquarema, small tributary of rio Fundo , about 1 km from town of Itapeba, near Maricá, 22°54’S 42°48’W . The following lots have only “general” locality information but are cited here because they were referred to by Schultz (1959: 112, USNM 94147 and USNM 177701) or were used by Myers to identify species in the aquarium trade and for publication of scientific names in Innes (1935 and later editions), USNM 94147 and USNM 129922 ; USNM 94147 , 3 , two females, one male, 29.4-40.5 mm SL, vicinity of Rio de Janeiro ; USNM 129922 , 1 , adult male, 32.2 mm SL ; USNM 177701 , 39 ,young to adults, 19.4-33.1 mm SL, Rio de Janeiro to Macaé ; USNM 249889 , 2 immatures, 28.5-34.7 mm SL, município de Nova Iguaçu, Tinguá, 22°41’S 43°25’W ; USNM 254256 , 10 , small adults to adults, 27.1- 34.2 mm SL ; MZUSP 19501 , 1 , adult male, SL 30.8 mm, município de Itaguaí, old road between cities of São Paulo and Rio de Janeiro at km 48 ; MZUSP 19334 , 5 , small adults to adults, 25.7-35.3 mm SL, município de Angra dos Reis, 23°00’S 44°19’W ; MNRJ 5570 , 40 , young to medium sized adults, 17.4-41.4 mm SL, município deAngra dos Reis, rio Ariró Pequeno ; USNM 249874 , 67 , young- small adults, 24.3-35.0 mm SL, município de Angra dos Reis, rio da Areia, first small tributary to rio Jurumirim on right side of road BR 155 , northeast of town of Angra dos Reis, 22°52’S 44°15’W ; USNM 249882 , 67 , young-adults, 21.2-46.3 mm SL ; MZUSP 26900 , 67 , young to small adults, 23.0- 31.2 mm SL. Following lots collected in São Paulo State. Município de Ubatuba : USNM 249883 , 48 immatures to large adults, SL 24.1-50.2 mm, rio Puruba , a small north branch about 5 km from road BR 101 , north and east of Ubatuba , 23°18’S, 44°56’W ; MZUSP 26901 , immatures-medium adults 23.9-33.5 mm SL . USNM 249872 , 15 , immatures, 21.6-29.7 mm SL, small tributary of rio Itamambuca , about 3 km from road BR 101 , 23°23’S 45°02’W ; MZUSP 26901 , immatures-medium adults 23.9-33.5 mm SL ; MZUSP 20462 , 5 , juveniles, 24.7-28.5 mm SL, valley between Jardim Acaraú and road BR 101 , approximately 23°26’S 45°05’W ; MZUSP 26896 , 9 , immatures to small adults, SL 20.4-31.5 mm ; MZUSP 26897 , 4 , young to small adult, 12.9-32.2 mm SL ; USNM 249873 , 25 immatures to adults, SL 25.0-47.0 mm SL, south branch of rio Tavares, rio Ubatuba system at Flora Brasilia Nursery, 23°27’S 45°05’W ; USNM 249876 , 7 immatures to small adults, 30.5-41.0 mm SL, rio do Cemitério, tributary to rio Ubatuba , 23°25’S 45°07’W ; USNM 313675 , 2 , adult-large adult, c&s, 1 female, 1 male, 37.2-57.3 mm SL ; USNM 249879 , 8 immatures to adults, 33.4-51.0 mm SL ; USNM 254261 , 8 , adults to large adults, 34.4-60.9 mm SL, rio Silva, a tributary to rio Ubatuba along road between Ubatuba and Taubaté, 23°23’S 45°07’W ; USNM 313671 , 2 , adult to large adults, c&s, 1 female, 1 male, 47.4-58.2 mm SL ; MZUSP 26899 , 10 , adults-large adults, 42.0- 61.0 mm SL ; USNM 249880 , adult to large adults 1 female, 1 male, 46.8-60.6 mm SL ; MZUSP 19820 , 02 , adults, 38.1-48.9 mm SL, ribeirão Ipiranguinha ; MZUSP 26909 , 2 , immature to small adult, 20.8-29.4 mm SL, stream on right side of the mouth of rio da Lagoa, Itaquá ; USNM 326812 , immature to adults 3, 36.0-48.0 mm SL, rio Indaiá, Fazenda Agricola Florestal Capricórnio Ltda. Município de Cananéia on road SP 193 km 56 .: USNM 326748 , immatures to adults 8, 28-43.3 mm SL ; MZUSP 53280 , immatures to adults 8, 31.0- 42.3 mm SL. Município de Caraguatatuba : USNM 249881 , 6 , small adults to medium adults, 31.7- 48.8 mm SL, rio Escuro , 23°28’S 45°12’W ; MZUSP 26906 , 8 , adults, 29.9 mm- 44.1 mm SL ; USNM 313684 , 6 , young to small adults, 24.2-32.1 mm SL, nameless stream 2.6 km up a dirt road from a point on road SP 55 17 km south of town of Ubatuba , 23°30’S 45°12’W ; MZUSP 53281 3, medium adults, 32.6- 40.7 mm SL, rio de Ouro, Reserva Florestal Município de Bertioga : USNM 362366 , 30 , young to adults, 21.3-45.7 mm SL, município de Bertioga, approximately 10 km north of town of Bertioga on road SP 98 between Bertioga and Mogi das Cruzes, an unnamed stream, 23°47’S 46°05’W ; MZUSP 53282 , 43 , young-adults, 21.7-38.8 mm SL ; FMNH 54887 , 1 , developing male, 37.8 mm SL. Município de Santos: Raiz da Serra , creek at base of mountains approximately 19 km from Santos, approximately 23°50’S 46°23’W ; CAS ( SU) 33869, 18, young to developing males, 17.4-31.4 mm SL; USNM 249887 , 2 , females, 39.4-48.8 mm SL, rio Biguá near Biguá, approximately 24°15’S 47°31’W ; USNM 249888 , adults to large adults, 37.5-57.3 mm SL ; USNM 254276 , 2 , adult female, adult male, 32.8-43.2 mm SL. Município de Iguape : ribeirão da Fazenda Cantagalo, Juquiá, approximately 24°19’S 47°36’W ; MZUSP 20218 , 6 , young to adults, 20.0- 46.3 mm SL ; MZUSP 53283 , 4 , adults, 34.2 mm, 48.3 mm SL, ribeirão da Fazenda Poço Grande, Juquiá ; USNM 254260 , 3 , adult males, 39.4-44.6 mm SL, Iguape, approximately 24°43’S 47°33’W . Following lots collected in Paraná State, município de Morretes (except where noted) . USNM 177820 , immatures 3, 23.5-27.8 mm SL, Morretes, on rio Marumbi, tributary of rio Nhundiaquara , 25°30’S 48°50’W . USNM 249897 , immatures to adults 4, 21.0- 46.5 mm SL, rio Nhundiaquara at Morretes, 25°29’S 48°49’W ; USNM 257114 , immatures to adults 6, 24.0-41.0 mm SL ; USNM 236089 , immature to adult 2, 34.3-45.8 mm SL, c&s ; USNM 257198 , immature, 29 mm SL and adult (tail removed for histology, SL not recorded) ; USNM 249886 , adults 2, 34.6-37.8 mm SL ; MZUSP 40281 , immatures to adults 28, 20.5-35.5 mm SL . MZUSP 53284 , immatures to adults 21, 16-46.8 mm SL, município Porto de Cima, rio São João at first large bridge east of São João on road between BR 116 northeast of Curitiba and Paranaguá, 25°23’S 48°50’W . MZUSP 53285 , immatures to adults, 22.5-43.0 mm SL, rio Sagrado east of Morretes , 25º32’S 48º46’W . Following lots collected rio Nhundiaquara , opposite and just downstream of Hotel Nhundiaquara , USNM 249890 , 6 adults, 32.5-48.5 mm SL ; USNM 249894 , adults 2, 38.5-44.8 mm SL ; USNM 257115 , adults 9, 30.8-49.8 mm SL. Following lots from state of Paraná State , MZUSP 17838 , immatures to adults 16, 15.3- 24.5 mm SL, rio Iraí, 6 km east of Curitiba ; USNM 270645 , immatures to adults 28, 20.0- 45.3 mm SL, rio Iraí, tributary of rio Iguaçu along road BR-425 east of Curitiba. Unnamed blackwater stream crossing road “ Elísio Pereira Alves Filho, about 15 km from road BR-227, southwest of Paranaguá , 25°40’S 48°33’W . USNM 306378 , immatures to adults, 62, 14.5-40.8 mm SL ; MZUSP 40228 , immatures to adults 62, 14.3-38.2 mm SL. Ribeirão Amola Faca, tributary to rio Iguaçu , near Balsa Nova in fazenda Lara Maria , USNM 326811 , 4 immatures to adults, SL 30.0- 46.5 mm SL Note: field photo made of largest specimen ; MZUSP 53286 , 6 adults, 38.0- 45.6 mm SL. Serrinha a small community between the towns of Contenda and Lapa, 1-2 km north of road BR-476, extending southwest from Curitiba . USNM 326810 , 72 immatures to adults, 21.7-43.0 mm SL ; MZUSP 53287 , 71 immatures to adults, 19.5-44.4 mm SL. Following lots collected in Santa Catarina State . MNRJ 6004 , immatures 13, 21.5-32.5 mm SL, unnamed tributary of rio Itapocuzinho , Guará Mirim ; MNRJ 3760 , immatures to adults 6, 37.0-55.0 mm SL, Joinville, date unknown. Rio Cubatão near Joinville , USNM 249893 , immatures to adults 25, SL 16.3-51.3 mm ; MZUSP 53288 , immatures to adults 10, 32.0- 45.0 mm SL. Rio Pique , Morro Cortado, off BR 101 at Meleiro near Limeira : USNM 254269 , 4 , 21.4-29.8 mm SL ; USNM 249892 , 10 , 17.6-24.3 mm SL ; MZUSP 19871 , 9 , 18.3-22.4 mm SL. Rio Jordão at Siderópolis, 28°38’S 49°27’W ; USNM 249899 , immatures to adults 24, 15.8-46.6 mm SL and MZUSP 26902 , immature to adults 24, 17.6-44.5 mm SL. Rio Faxinalzinho at Mãe dos Homens, near Praia Grande , approximately 29°15’S 50°06’W , USNM 279877 , 1 adult 1, 44.0 mm SL ; MZUSP 53289 , immature 1, 32.2 mm SL ; MCP 13766 , immature 1, 31.0 mm SL. Arroio Lindo on road BR-101, unnamed tributary of rio Cubatão , Joinville : USNM 326737 , immatures to adults 3, 32.7-44.2 mm SL ; USNM 326738 , immature 1, 35.0 mm SL ; MZUSP 53290 , immatures to adults 3, 30.0- 45.8 mm ; MCP 21698 , immatures 2, 36.4-39.0 mm SL. Unnamed stream tributary to rio Itapocu at Corupá : USNM 279876 , immatures to adults 4, 34.3- 51.0 mm SL ; MZUSP 53291 , immatures to adults 4, 27.5-48.3 mm SL ; MCP 216699 , immatures to adults 3, SL 22.0- 46.3 mm SL Município de Praia Grande, a tributary of rio Grande about 3 km from Praia Grande : USNM 362362 , immatures to adults 10, 27.5- 42.7 mm SL ; MZUSP 53292 , immatures to adults 10, 28.8-38.0 mm SL. Município de Praia Grande 29º14’S 50º01’W , rio Canoas, 8 km from Praia Grande : USNM 326747 , 42 immatures to adults, 27.8- 45.0 mm SL ,; MZUSP 53293 , immatures to adults, 42, 24.0 42.5 mm SL ; MCP 21701 , immatures to adults 42, 25.3-48.0 mm SL; Rio Canoas 13 km above Praia Grande : USNM 326743 , immatures 4, 30.0- 37.2 mm SL ; MZUSP 53294 , immatures 3, SL 30.0-36.0 mm ; MCP 21699 , immatures 3, 29.8-33.5 mm SL. Following lots collected in Rio Grande do Sul State . MCP 10145 , immatures 10, 16.3-20.0 mm SL, from lagoa Emboaba, Tramandaí . MAPA 2126 , immatures to adult 8, 18.2-35.7 mm SL, from arroio Água Parada, Maquiné,tributary to rio Maquiné near town of Maquiné approximately 29º40’S 50º12’W : USNM 326746 , immatures-adults 5, SL 27.5-45.3 mm SL ; MZUSP 53295 , immatures to adults 4, 31.0-41.0 mm SL ; MCP 21702 , immatures to adults 4, 30.8-41.5 mm SL. Rio Pinheiro above Maquiné : USNM 326745 , immatures to adults 9, 28.8-49.5 mm SL ; MZUSP 53296 , immatures to adults 9, SL 33.4-54.2 mm ; MCP 21703 , immatures to adults 8, 33.0-52.0 mm SL, arroio Sanga do Icó tributary of rio Maquiné, about 2 km from town of Barra do Ouro ,: USNM 326744 , immatures to adults 9, 26.5-43.0 mm SL ; MZUSP 53297 , immatures to adults 9, 23.5-39.5 mm SL ; MCP 21704 , immatures to adults 9, 27.0- 35.5 mm SL. Rio Três Pinheiros 8 km from road BR-101 near Itati 29º32’S 50º06’W , rio Três Forquilhas drainage : USNM 326740 , immatures 3, 16.0-34.0 mm SL ; MZUSP 53298 , immatures 4, SL 14.7-30.0 mm ; MCP 21705 , immatures 4, 17.0- 35.2 mm SL. Rio Mitmann in Vila Nova, about 10 km from road BR-101, rio Três Forquilhas drainage : USNM 326739 , immatures-adults 21, 25.5-45.3 mm SL ; MZUSP 53299 , immatures to adults 20, 25.0- 43.3 mm SL ; MCP 21706 , immatures to adults 21, 25.7-45.5 mm SL. município de Três Cachoeiras, rio das Pacas near Morro Azul 29º24’S 49º57’W : USNM 326736 , immatures-adults 25, 15.7-33.5 mm SL ; MZUSP 53300 , immatures to adults 26, 19.0- 35.6 mm SL ; MCP 21707 , immatures to adults 26, 16.8-36.0 mm SL. Município de Torres , rio dos Mengue between Morro Azul and Rua Nova 29º17’S 49º55’W : USNM 326742 , immatures to adults 9, 24.5-42.5 mm SL ; MZUSP 53301 , immatures to adults 9, 20.5-36.7 mm SL ; MCP 21708 , immatures to adults 9, 20.3-38.0 mm SL. Canal between lagoa da Emboaba and lagoa Emboabinha on road between Tramandaí and Osório : USNM 326741 , immatures-adults 27, 16.3-33.5 mm SL ; MZUSP 53302 , immatures to adults 24, 17.5-35.6 mm SL ; MCP 21708 , immatures to adults 24, 17.5- 41.3 mm SL .

Figures

Fig. 77. Mimagoniates microlepis, young adults male below, female above, USNM 254261, 61.2.0 mm and 43.0 mm SL, respectively, Brazil, São Paulo State, rio Silva on road between Ubatuba and Taubaté.

Fig. 78. Mimagoniates microlepis, large adult male, USNM 249888, 53.3 mm SL, Brazil, São Paulo State, rio Biguá near Biguá.

Fig. 79. Mimagoniates microlepis, adult male, MZUSP 14668, 40.7 mm SL, Rio de Janeiro State, rio Guapiaçu, Cachoeiras de Macacu.

Fig. 81. Mimagoniates microlepis, large adult male, USNM 249876, 59.3mm SL; anterior 13 anal-fin rays, lateral view, left side.

Fig. 83. Mimagoniates microlepis, large adult male, USNM 249876, 59.3 mm SL; pelvic-fin rays, ventral view, left side.

Fig. 84. Mimagoniates microlepis, adult male, USNM 249885, 41.2 mm SL; pelvic-fin rays, ventral view, left side. All rays with fewer branches compared to those of large adult male in Fig.83.

Fig. 85. Mimagoniates microlepis, adult male, USNM 236089, 45.3 mm SL; osteology of caudal skeleton and caudal-fin squamation (lower left), lateral view, left side. Principal caudal ray 12 indicated. Relationship of modified dorsal lobe caudal-fin squamation to caudal gland shown in inset at lower left. Distal regions of ventral and posterior scales and their epidermis form flap partly covering intake openings of pump chamber. Modified from Menezes et al. (1988).

Fig. 86. Mimagoniates microlepis, adult male, USNM 249876, 59.3 mm SL; jawsand dentition, lateral view, right side, anterior atright.Comparenumber ofmaxillary teeth(10)in this specimen with that in Fig. 87 (6). Number of maxillary teeth varies considerably in specimens depending on their standard length.

Fig. 88. Mimagoniates microlepis. Dorsal-fin height as function of SL by sex.

Fig. 89. Comparative plots of the number of branched anal-fin rays for population samples of Mimagoniates microlepis within range of species expressed by Tukey box plots based on percentiles.

Fig. 90. Comparative plots of (a) Number of pored lateral-line scales; (b) Number of horizontal scale rows on body; (c) Number of lateral-series scales; (d) Number of horizontal scale rows around the caudal peduncle; (e) Number of predorsal scales; (f) Total number of vertebrae for samples of Mimagoniates microlepis within range of species expressed by Tukey box plots. For I, II, III, IV and V locations, see full legends at Fig. 89.