A M ERIC AN MUSEUM NOVITATES Number 3831, 18 pp. April 21, 2015 Phylogenetics of Teleogramma, a riverine clade of African cichlid fishes, with a description of the deepwater molluskivore—Teleogramma obamaorum— from the lower reaches of the middle Congo River MelAnie l.J. StiASSny1 And S. elizAbeth AlteR2 AbStRACt The lower Congo River and nearby habitats harbor numerous endemic lineages of cichlid fishes, including some with highly specialized morphologies. based on morphological and molecular data, we herein describe a new species of Teleogramma, a member of the chromi- dotilapiine clade found on rocky outcrops in the lower reaches of the middle Congo River. The new species, T. obamaorum, is distinguished from congeners by numerous morphological and ecological attributes, including the lack of dorsoventral head and body depression, absence of sexual dichromatism, and features of laterosensory anatomy, pharyngeal and gut morphology, and dietary preference. Phylogenetic analyses of two nuclear and two mitochondrial loci using bayesian and maximum-likelihood inference lend strong support for the taxonomic validity of T. obamaorum and provide preliminary estimates of species relationships within the genus. The discovery of a new, ecomorphologically distinctive cichlid species in the Congo River suggests that additional research focus on riverine clades has the potential to greatly contribute to our understanding of evolutionary dynamics in this hyperdiverse group of teleost fishes. Keywords: Afrotropics, riverine biodiversity, molecular systematics, taxonomy 1 department of ichthyology, division of Vertebrate zoology, American Museum of natural history. 2 department of biology, york College, City University of new york. Copyright © American Museum of Natural History 2015 ISSN 0003-0082 2 AMeRiCAn MUSeUM noVitAteS no. 3831 intRodUCtion Teleogramma is a small clade of rheophilic, rock-dwelling cichlids restricted to fast-flowing waters in the western Congo basin. Unlike members of most riverine cichlid lineages, which exhibit muted morphological innovation and share a rather generalized “riverine cichlid” gestalt, Teleogramma species have highly specialized anatomies. When first described by bou- lenger (1899), the genus was erroneously placed in the family labridae. Myers (1939) and subsequent studies placed it within the Cichlidae, and all noted the highly distinctive nature of these unusual riverine cichlids. takahashi and nakaya (2001) provided a detailed anatomical description of Teleogramma and enumerated a suite of osteological and myological synapomor- phies for the genus. Species within this genus are readily recognized by their markedly dorso- ventrally compressed heads bearing a pair of prominent tubular nostrils, their elongate compressed bodies, and the presence of a single continuous lateral line not interrupted into upper and lower branches, as in most other cichlids. due in part to their highly autapomorphic morphology, determination of the interrelation- ships of Teleogramma within the Cichlidae has proven elusive in morphological analyses (e.g., Stiassny, 1997; takahashi and nakaya, 2001). however, a recent multilocus phylogeny (Schwar- zer et al., 2014) provides conclusive support for the placement of Teleogramma as nested within the chromidotilapiine lineage of African cichlids. Chromidotilapiines are a primarily riverine clade that originated in West and Central Africa in the late eocene/early oligocene (Schwarzer et al., 2014), and therefore represent an early African cichlid radiation with origins long predat- ing those of the hyperdiverse, predominately lacustrine, assemblages of eastern Africa. Prior to the present study, Teleogramma was considered to contain four species (Roberts and Stewart, 1976), three of which are endemic to rocky outcrops in fast-flowing waters along stretches of the lower Congo River (lCR): T. brichardi (fig. 1A), T. gracile (fig. 1b), and T. depressa (fig. 1C). A fourth species, T. monogramma (fig. 1d), occurs in rapids-associated outcrops below the confluence of two Congo tributaries, the lulua and Kasai, located some 850 river kilometers upstream from the outflow of the lCR from Pool Malebo. during a period of exceptionally low water throughout western Congo in the summer of 2011, a series of highly distinctive specimens were collected in newly exposed rocky outcrops along the lower reaches of the middle Congo River just upstream of Pool Malebo (fig. 2). These specimens were unassignable to any previously described species. Preliminary morphological examination suggested that, although lacking a number of distinctive features of the genus described by takahashi and nakaya (2001), these specimens likely represented an undescribed species of Teleogramma, a finding fully corroborated in the present study with both morpho- logical and molecular evidence. in addition to detailed morphological examination, and in order to phylogenetically place the new species within the genus, representatives of all putative Teleogramma species and a chro- midotilapiine outgroup were sequenced for two nuclear (Sh3PX3 and Ptr) and two mitochon- drial (Co1 and nd2) loci. Model-based phylogenetic reconstructions were undertaken on these molecular data and the results of those analyses provide a comparative framework for the formal taxonomic description of the distinctive new species, Teleogramma obamaorum, provided herein. 2015 StiASSny And AlteR: PhyloGenetiCS oF TELEOGRAMMA 3 FiGURe 1. Teleogramma (boulenger, 1899) general habitus of currently recognized species, in lateral aspect, ♂ above, ♀ below. A, T. brichardi (AMnh 246986); B, T. gracile (AMnh 247261); C, T. depressa (AMnh 239325); D, T. monogramma (AMnh 247806 ♂, 253127 ♀). Scale bars = 1 cm. Materials and Methods Morphology: Fourteen standard morphometric measurements and seven meristic counts were taken following barel et al. (1977). Specimens were carefully pinned flat and photographed on the left side using a nikon digital SlR camera with a 60 mm f/2.8 AF Micro-nikkor lens. linear measurements were then taken using the open access software imageJ v1.48 (Schneider et al., 2012), and vertebral and fin-ray counts were taken from radio- graphed and/or cleared and stained specimens. Gill raker counts are totals for the first arch and include the raker at the junction of ceratobranchial and epibranchial elements. lateral- line counts exclude the small pored scales on the caudal fin distal to the point of caudal flexion. For comparative purposes, corresponding counts and measurements for the closely related T. monogramma and T. brichardi are provided. Additional specimens of all putative species were examined and cleared and stained using a modified protocol of taylor and Van dyke (1985). Abbreviations used throughout the text are: C&S, cleared and stained preparations; SL, standard length; HL, head length; ex, number of specimens examined. institutional abbrevia- tions follow Sabaj Perez (2014). Molecular Methods: total genomic dnA was extracted from fin clips using the dneasy tissue extraction kit (Qiagen). Partial sequence fragments of four genetic markers (two mitochon- drial: cytochrome oxidase i (Coi) and nAdh dehydrogenase subunit 2 (nd2) (Kocher et al., 1995), and two nuclear: Sh3PX3 and Ptr (li et al., 2007)) were sequenced across representatives of all putative Teleogramma species. based on the phylogenetic hypothesis of Schwarzer et al. (2014), the type species of the genus Chromidotilapia sensu stricto (Chromidotilapia kingsleyae) was selected as an outgroup for rooting the Teleogramma trees. dnA amplification via PCR was performed with illustra PuRetaq Ready-to-Go PCR beads (Ge healthcare). For Coi, we used primers VF2_t1, 5′-tGtAAAACGACGGCCAGtCAACCAACCACAAAGACAttGGCAC-3′, FishF2_t1, 5′-tGtAAAACGACGGCCAGtCGACtAAtCAtAAAGAtAtCGGCAC-3′, 4 AMeRiCAn MUSeUM noVitAteS no. 3831 FishR2_t1, 5′-CAGGAAACAGCtAtGACACttCAGGGtGACCGAAGAAtCAGAA-3′, and FR1d_t1, 5′-CAGGAAACAGCtAtGACACCtCAGGGtGtCCGAARAAyCARAA-3′ (ivanova et al., 2007) with the following amplification conditions: 94° C for 2 min, 35 cycles of 94° C for 30 s, 52° C for 40 s, and 72° C for 1 min, with a final extension at 72° C for 10 min. For the nd2 marker, we used primers nd2trp, 5′-CAtACCCCAAACAtGttGGt-3′ and nd2Met, 5′-GtSGSttttCACtCCCGCttA-3′ (Kocher et al., 1995) with the following amplification conditions: 94° C for 2 min, 35 cycles of 94° C for 30 s, 55° C for 30 s, and 72° C for 1 min, with a final extension at 72° C for 10 min. For nuclear markers, amplifications were performed using primers Ptr_F458, 5′-AGAAtGGAtWACCAACACytACG-3′ and Ptr_R1248, 5′-tAAGGCA- CAGGAttGAGAtGCt-3′, and Sh3PX3_F461 (5′-GtAtGGtSGGCAGGAACytGAA-3′) and Sh3PX3_R982 (5′-CAAACAKCtCyCCGAtGttCtC-3′) (li et al., 2007), with the following amplification conditions for both: 94° C for 2 min, 35 cycles of 94° C for 30 s, 55° C for 40 s, and 72° C for 1 min, with a final extension at 72° C for 10 min. Successful amplifications were sequenced at Genewiz inc. (South Plainfield, nJ). Forward and reverse chromatograms were edited and aligned using Geneious R7.0 (bio- matters ltd., Aukland, nz). Alignment across all specimens was performed using MUSCle v3.5 (edgar, 2004), and best-fit models of nucleotide evolution were determined using JModelt- est (evolutionary biology Centre, Uppsala University, Sweden). to accommodate heterogeneity between gene regions, model-based analyses were conducted on the concatenated alignment partitioned into gene regions with parameters unlinked. likelihood analyses were carried out in RAxMl v7.2.8 black box (Stamatakis, 2006), and bayesian phylogenetic inference using Markov Chain Monte Carlo (MCMC) methods was implemented in Mrbayes v3.2 (huelsen- beck and Ronquist, 2001). Runs were performed using 15 million generations with trees sam- pled every 1000 generations, and a burn-in of 25%. in the likelihood analyses nodal support is given by means of bootstrap character resampling (1000 pseudoreplicates), and bayesian nodal support was assessed using Markov chain Monte Carlo sampling to derive posterior probabili- ties. Specimen voucher data and Genbank accession numbers for all loci generated in the current study are provided in table 1. ReSUltS Phylogenetic Analyses likelihood and bayesian model-based phylogenetic analyses resulted in the same topology (fig. 3), and provide strong support for the generic placement and taxonomic validity of Teleo- gramma obamaorum. The results of this small-scale analysis also provide a preliminary estimate of species relationships with the genus, which is partitioned into two main clades. Within “clade A” the new species, T. obamaorum, is resolved as sister to the Kasai/lulua–dwelling T. monogramma, and together these are sister to T. brichardi. We note that T. brichardi, as recog- nized herein, is restricted to specimens from the immediate vicinity of the type locality at the fishing village of Kinsuka, located on the left bank of the lCR at the first rapids downstream of Pool Malebo in the democratic Republic of Congo (fig. 2). 2015 StiASSny And AlteR: PhyloGenetiCS oF TELEOGRAMMA 5 s. e s 5 3 2 1 0 4 6 5 7 9 7 8 9 0 4 3 8 1 2 y 1 1 1 1 1 0 0 0 0 0 9 0 9 0 1 0 9 0 0 the anal sh3px3   KP7142   KP7142 KP7142 KP7142 KP7142 KP7142 KP7142 KP7142 KP7142 KP7142 KP7141 KP7142 KP7141 KP7142 KP7142 n/a KP7142 n/a KP7141 KP7142 KP7142 n d i ber 6 1 4 3 2 4 6 5 8 0 8 9 0 1 5 7 9 2 3 e m 9 9 9 9 9 8 8 8 8 9 7 8 8 8 9 8 7 8 8 nclud n Nu r P7141 P7141 P7141 P7141 P7141 P7141 P7141 P7141 P7141 P7141 P7141 P7141 P7141 P7141 P7141 a P7141 a P7141 P7141 P7141 ces i essio pt   K   K K K K K K K K K K K K K K n/ K n/ K K K n c ue Ac 8 6 5 9 1 0 2 4 3 3 5 6 7 1 4 7 8 gene seq GenBank co1   KP71415   n/a KP71415 KP71415 n/a KP71414 KP71415 KP71415 KP71415 KP71415 KP71414 KP71415 KP71414 KP71414 KP71415 n/a KP71414 n/a KP71414 KP71414 KP71414 e h mbers for t nd2   KP714163   KP714162 KP714165 KP714170 KP714177 KP714174 KP714159 KP714169 KP714160 KP714176 KP714167 KP714161 KP714168 KP714164 KP714171 hM101360 KP714175 hM101359 KP714172 KP714166 KP714173 u n ank accession Tissue code   227313   224324 223886 223885 223881 055-5455 055-5483 055-5482 061-6024 061-6026 024-2338 061-6025 035-3425 035-3426 224163 060-5942 060-5945 060-5943 033-3212 043-4296 043-4297 b mbers, and Gen AMNH catalog AMnh 263142 AMnh 255206 AMnh 255329 AMnh 255329 AMnh 255329 AMnh 247831 AMnh 247780 AMnh 247780 AMnh 247819 AMnh 246986 AMnh 240012 AMnh 246986 AMnh 240013 AMnh 240013 AMnh 255381 AMnh 239324 AMnh 239324 AMnh 239324 AMnh 241068 AMnh 246477 AMnh 246477 u     n g o p. p. p. p. xa, voucher catal Taxon P apia kingsleyae a obamaorum, n. s a obamaorum, n. s a obamaorum, n. s a obamaorum, n. s a monogramma a monogramma a monogramma a monogramma a brichardi a brichardi a brichardi a depressa a depressa a depressa a depressa a depressa a depressa a depressa a gracile a gracile table 1. ta oUtGRoU Chromidotil inGRoUP Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm Teleogramm 6 AMeRiCAn MUSeUM noVitAteS no. 3831 FiGURe 2. Collection sites of T. obamaorum, new species (red stars) along lower reach of middle Congo above Pool Malebo. A, exposed rocks in midstream of Congo River main channel just above Pool Malebo; B, collection site of holotype, among exposed rocks extending into main channel. inset photographs taken in August 2011 at a time of record low water throughout western Congo. While support is strong for the monophyly of “clade b” Teleogramma, which comprises the remainder of the lCR populations, branch lengths in this region of the tree are extremely short and the resultant topology highlights conflict between morphology-based taxonomy versus molecular signal. Such short branch lengths, in conjunction with molecular and morphological discord, suggest that rapid and recent divergence, and perhaps ongoing gene flow between incipient and/or young species, may be confounding the molecular signal provided by the four molecular markers used in this study. to further investigate the popu- lation dynamics, relationships, taxonomic status, and age of the lCR Teleogramma we are currently analyzing a genomewide sampling of Teleogramma populations utilizing restric- tion site associated (ddRAd) markers (Alter et al., in prep.). despite the inability of our four-locus dataset to entirely resolve the species composition and relationships among the lCR “clade b” Teleogramma, in contrast “clade A” taxa are well supported by these data. based on our molecular analyses, and a series of morphological and ecological attributes discussed below, we provide a formal taxonomic description of a new species of Teleo- gramma, T. obamaorum, a taxon geographically located between the widely disjunct lCR and Kasai/lulua populations. 2015 StiASSny And AlteR: PhyloGenetiCS oF TELEOGRAMMA 7 Chromidotilapia kingsleyae T. depressa (upstream Luozi) T. gracile (Lenga Lenga) T. gracile (Lenga lenga) C T. depressa (Inga) L A T. depressa (Inga) D 9999 T. depressa (Inga) E 1 T. depressa (Foulakari) B T. depressa (Mbudi) T. depressa (Foulakari) T. brichardi (Kinsuka) 92 T. brichardi (Kinsuka) 1 T. brichardi (Kinsuka) T. monogramma (Bukonde, R. Lulua) C 100 T. monogramma (Nsanga, R. Lulua) L .96 1 T. monogramma (Nsanga, R. Lulua) AD T. monogramma (Katende, R. Lulua) E A .99 T. obamaorum (Nkana) T. obamaorum (Maluku) 100 0.02 1 T. obamaorum (Maluku) T. obamaorum (Maluku) FiGURe 3. Maximum-likelihood phylogeny of Teleogramma (baysian analysis resulted in an identical topol- ogy). bootstrap support (>90%) indicated above internal nodes and posterior probabilities (>0.90) below. Collection localities of sampled individuals indicated in parentheses. Teleogramma obamaorum, new species Figures 4–8, tables 2 and 3 holotype: AMnh 264012, male, 75.00 mm Sl, democratic Republic of Congo, Kinshasa Province, main channel of middle Congo River at rocky outcrop en route to Mayi ndombe River (4° 01′ 31.4″S, 15° 36′ 37.1″e), M.l.J. Stiassny et al., 10 August 2011. Paratypes: AMnh 254857, 6 paratypes, 44.6–63.4 mm Sl, same locality as holotype. — MRAC b5-04-P-1, 1 paratype, 59.4 mm Sl, same locality as holotype. — AMnh 255206, 6 paratypes, 52.1–68.2 mm Sl, democratic Republic of Congo, Kinshasa Province, main channel of middle Congo River at rocky outcrop above guard post at nkana (3° 53′ 31.4″S, 15° 55′ 27.9″e), M.l.J. Stiassny et al., 14 August 2011. — MCz 171655, 1 paratype, 63.0 mm Sl, demo- cratic Republic of Congo, Kinshasa Province, main channel of middle Congo River at rocky outcrop above guard post at nkana (3° 53′ 31.4″S, 15° 55′ 27.9″e), M.l.J. Stiassny et al., 14 August 2011. — AMnh 255329, 4 paratypes, 48.8–64.9 mm Sl, democratic Republic of Congo, Kinshasa Province, main channel of middle Congo River at rocky outcrop above Maluku (4° 00′ 48.6″S, 15° 38′ 42.8″e), M.l.J. Stiassny et al., 14 August 2011. —zSM 43801, 1 paratype, 58.9 mm Sl, democratic Republic of Congo, Kinshasa Province, main channel of middle Congo River at rocky outcrop above Maluku (4° 00′ 48.6″S, 15° 38′ 42.8″e), M.l.J. Stiassny et al., 14 August 2011. 8 AMeRiCAn MUSeUM noVitAteS no. 3831 FiGURe 4. T. obamaorum, new species, general habitus, lateral view, in preservation. A, ♂ holotype (AMnh 264012); B, ♀ paratype (AMnh 254857); C, schematic representation of genital papillae. Scale bar = 1 cm. Additional nontype Material examined: AMnh 264013, 3 specimens, democratic Republic of Congo, Kinshasa Province, main channel of middle Congo River at rocky outcrop en route to Mayi ndombe River (4° 01′ 31.4″S, 15° 36′ 37.1″e), M.l.J. Stiassny et al., 10 August 2011. — AMnh 264014, 3 specimens, democratic Republic of Congo, Kinshasa Province, main channel of middle Congo River at rocky outcrop above guard post at nkana (3° 53′ 31.4″S, 15° 55′ 27.9″e), M.l.J. Stiassny et al., 14 August 2011. — AMnh 264015, 3 specimens, 2 C&S, democratic Republic of Congo, Kinshasa Province, main channel of middle Congo River at rocky outcrop above Maluku (4° 00′ 48.6″S, 15° 38′ 42.8″e), M.l.J. Stiassny et al., 14 August 2011. diagnosis: Teleogramma obamaorum is distinguished from all congeners by the absence of a sexual dichromatism of the caudal fin (versus caudal fin of females with a distinctively marked dorsal blaze), markedly inflated pores in the laterosensory system of the head, jaws, suspensorium and infraorbital series. it has 5 separate infraorbital elements (versus 4 in con- geners), and a robust lower pharyngeal jaw (versus a markedly more gracile element in conge- ners). Uniquely among congeners, T. obamaorum is a robust species lacking dorsoventral head and body depression. it is further readily distinguished from T. monogramma by number of anal spines (7–8 versus 4), and from T. brichardi in the possession of 32–35 pored lateral-line scales (versus 51–60). description: A Teleogramma species attaining maximum-recorded size of 75.0 mm Sl (mature male holotype, AMnh 264012), with general body shape and appearance in figure 4. Proportional measurements for holotype and 19 paratypes, and comparable ranges for T. mono- gramma and T. brichardi, the taxa that molecular analysis suggests most closely related (fig. 3), 2015 StiASSny And AlteR: PhyloGenetiCS oF TELEOGRAMMA 9 FiGURe 5. T. obamaorum, new species. A, oral jaws and suspensorium; B, first gill arch; C, infraorbital series; D, T. monogramma, infraorbital series; E, T. brichardi, infraorbital series. are given in table 2 and meristic attributes summarized in table 3. body deep and laterally compressed, with greatest depth at level of first dorsal fin spine. dorsal and ventral body pro- files slightly convex to relatively short, deep, caudal peduncle. head robust rather than dorso- ventrally compressed. Adductor mandibulae muscle complex markedly enlarged lending bulging appearance to the cheeks. Snout prominent and bears single, tubular, nostril on either side. Snout and dorsal head profile rise at angle of 35°–40° to midorbit, often with small, fat- filled, nuchal hump just anterior to dorsal fin origin in both sexes. dorsal fin XXi–XXiii (mode: XXii) 8 or 9 (mode: 9). Anal fin Vii–Viii (mode: Vii) 8–9 (mode 9); both with prominent, scaleless, fleshy bases. dorsal-fin spines gradually increase in length posteriorly to around 8th spine then coequal in length. Posteriormost branched fin rays of dorsal and anal fins reach to anterior third of caudal fin. Caudal fin large and paddle shaped, with 14-branched rays (7+7) and rounded symmetrical distal margin. Pectoral fins broadly fan shaped, long but falling short of anus, with 15–16 rays. Pelvic fins falling just short of anus, with third branched ray longest in both sexes. Jaws prominent, slightly prognathous and robust with lips well developed and fleshy. inner and outer row teeth of both jaws pointed unicuspid. Single series of 2–3 enlarged, recurved canines situated anteriorly on premaxilla. dentary with 3 enlarged canines anteriorly, larger pair slightly displaced dorsolaterally. Remaining outer row of teeth in both jaws gradually taper in size and extend almost entire length of both jaws (fig. 5A). Four or 5 inner rows of small, slightly recurved, unicuspid teeth clustered on anterior third of dentary; no inner rows distally on lower jaw. two or three inner tooth rows anteriorly on premaxilla, tapering to single row terminating at two-thirds of length of dentigerous arm. lower pharyngeal jaw is markedly more robust than in congeners with slightly sinuous, rather than straight, ventral suture (com- pare e.g., figs. 6A and b, C). Usually 15–16 moderately robust, unicuspid teeth in posterior row. Symphysial teeth enlarged and somewhat molariform (fig. 6A). Gill rakers elongate, nonden- ticulate and slender (fig. 5b). eight to 10 rakers along outer row of first gill arch; 5–6 cerato- branchial rakers, usually one raker in angle of arch, and 2 or 3 epibranchial rakers. 10 AMeRiCAn MUSeUM noVitAteS no. 3831 - o n 4 7 8 3 2 0 8 2 8 7 2 7 5 mo 0) SD 1.2 1.2 0.8 1.2 0.6 0.9 1.5 1.1 3.7 1.8 1.0 1.8 2.8 2 e ranges for T. T. brichardi (n = MeanRange 84.347.8–87.7 16.815.1–19.9 28.525.0–30.3 27.426.1–28.8 61.459.1–63.5 10.49.2–11.4 16.314.6–17.4 23.921.2–26.8 66.764.2–68.9 45.141.3–53.0 30.026.5–33.6 21.620.0–24.0 30.026.5–33.6 16.112.0–21.8 v ati r a p m 7 7 4 0 6 7 8 7 1 8 3 6 3 co 20) SD 1.4 1.2 0.9 1.1 0.4 1.0 1.2 1.7 3.3 1.6 1.7 2.5 1.4 s with ma (n = e 75.4 16.7 28.0 29.1 63.8 0.0 19.9 25.7 67.4 57.3 39.1 26.4 34.0 17.9 pecie gram Rang 49.9– 12.0– 23.3– 25.9– 59.6– 85.–1 15.8– 20.6– 61.3– 42.7– 32.4– 19.2– 23.1– 13.2– w s ono n m, ne T. m Mea 60.6 14.7 26.6 27.2 61.0 9.2 17.9 23.0 64.7 46.9 36.3 21.9 28.1 14.8 u r o a m 8 1 0 0 8 1 2 5 5 7 0 0 7 oba es SD 0.8 1.5 1.3 1.9 0.3 1.0 1.7 1.8 4.4 2.2 1.5 2.2 2.3 mma speci e 75.0 21.8 31.2 32.3 64.2 1.2 16.1 31.6 66.5 55.3 39.5 24.2 34.5 19.6 Teleogra ama new nRang 44.6– 18.4– 25.8– 27.6– 57.6– 9.5–1 12.0– 24.7– 60.2– 39.4– 31.4– 19.6– 26.0– 10.7– s of ma ob Mea 20.3 29.0 30.2 60.6 10.3 14.2 28.1 64.1 45.3 35.7 21.7 29.9 14.5 e eri am N 20 20 20 20 20 20 20 20 20 20 20 20 20 e s ogr p e e ty Tel yp a for Holt 75.0 21.8 27.5 29.5 60.8 10.5 14.8 29.6 64.5 55.3 38.3 21.2 29.0 15.0 at d c di. table 2. Morphometrigramma and T. brichar Standard length (mm) % Standard length body depth head length Predorsal length Preanal length Caudal peduncle depth Caudal peduncle length Anal-fin base length dorsal-fin base length % Head Length lower-jaw length Upper-jaw length eye diameter Snout length interorbital width