Dorsal Fin Spines Research Articles

A partial articulated acanthodian from the Silurian of New Brunswick, Canada

Vertebrates preserved in calcareous concretions collected from Silurian marginal-marine deposits near Nerepis, southern New Brunswick include agnathan thelodonts, a heterostracan, and possibly an anaspid, as well as spines and scales from an acanthodian (the only jawed fish remains). Several incomplete specimens of acanthodians, all determined as the same taxon Nerepisacanthus denisoni nov. gen. et sp., are described based on material identified in several museum collections. The fish-bearing nodules are from the Silurian Cunningham Creek Formation, which has been dated from ?late Llandovery (Telychian) – Ludlow (Ludlovian). The new species is only the second undisputed pre-Devonian acanthodian taxon known from articulated, although incomplete, specimens. N. denisoni has robust flank scales with a smooth crown bearing short subparallel ridges along the anterior edge, plus flatter scales with a crown formed of elongate, curved, areal-growth ridges, anterior to the pectoral region. The anterior dorsal fin spine is long, slightly curved and bears up to seven smooth longitudinal ridges of varying width on each side. The perichondrally ossified scapula is thin-walled and probably has a U-shaped cross section. “Dentition” cones and small denticles are preserved in the branchial or pharyngeal region. Although elements indisputably identifiable as dentigerous jaw bones were not found in the specimens, N. denisoni appears most closely related to the Early Devonian ischnacanthiform Acritolepis and is tentatively assigned to the family Acritolepidae Valiukevičius and Burrow.

Genetic and morphological divergence within the Sebastes pachycephalus complex (Scorpaeniformes: Scorpaenidae)

Genetic and morphological divergence among the four subspecies in the Sebastes pachycephalus complex (S. pachycephalus pachycephalus, S. p. nigricans, S. p.nudus and S. p. chalcogrammus) was clarified. Principal coordinate analysis (PCoA) based on AFLP clearly divided 55 specimens of the complex into two groups, the S. p. pachycephalus–S. p. nigricans group (P-Ni group) and the S. p. nudus–S. p. chalcogrammus group (Nu-C group), although three specimens occupied intermediate positions. The minimum spanning network (MSN) based on partial sequences of the mitochondrial control region (mtCR) failed to separate either the P-Ni and Nu-C groups or the four subspecies into distinct clades, although restricted gene flow and genetic differentiation between the former were indicated by the FST estimation. Differences in morphological characters, including counts of pectoral fin rays and counts of dorsal fin spines lacking basal scales, were also evident between the two groups. However, little or no genetic or morphological difference was found between the two subspecies within each group. It was concluded that the P-Ni and Nu-C groups of the S. pachycephalus complex actually represent two different species, which is further supported by their sympatric distribution. Differences in dorsal body coloration and the presence or absence of brown spots on the ventral surface, which were formerly used to discriminate between four “subspecies,” may simply represent intraspecific variation. The three specimens occupying intermediate positions in the AFLP PCoA also occupied equivocal positions between the two species in the principal component analysis (PCA) based on morphometric characters, suggesting that they were hybrids between the two species. The star-shaped MSN of mtCR, which lacks distinct clades representing the two species, may be due to not only interspecific hybridization but also the sharing of ancestral haplotypes.

Spinipterus, a new genus of small, spiny catfish (Siluriformes: Auchenipteridae) from the Peruvian Amazon

A new species of small auchenipterid catfish is described based on a single specimen from the Amazon basin and is placed into a new genus. Spinipterus, new genus, is diagnosed by the following striking autapomorphic features: pectoral and dorsal-fin spines with four prominent rows of serrations; lateral margin of the skull roofing bones ornamented with a single row of spines; size reduction of hipurals; elongation and angle of posterior cleithral process; and adducted dorsal fin rests into a mid-dorsal groove. The new genus also differs from most other auchenipterid genera by having lower counts for some fin rays. Current diagnoses of auchenipterid genera do not allow us to place the new species into any genus, which requires us to propose the new genus Spinipterus.

Age and growth of the sailfish Istiophorus platypterus (Istiophoridae) in the Gulf of Tehuantepec, Mexico

We analysed the recreational-dependent and the commercial fishery data of Istiophorus platypterus from the Gulf of Tehuantepec, Mexico. We estimated the age and growth of sailfish in our study based on growth rings in the cross-sections of the fourth dorsal fin spine. We sampled 4976 individuals, with length and weight data collected from 2000 to 2008. We analysed the ages of 535 sailfish, with their individual growth estimated by using two models. The first was the Schnute generalized-growth model and the second was the von Bertalanffy growth function. The parameters in each model and their confidence intervals (CI) were computed. The best candidate model was selected using Akaike's information criterion. We found the growth curve for sailfish from the Gulf of Tehuantepec is based on the three-parameter model rather than the generalized Schnute model. The parameters estimated and confidence intervals were (1) mean asymptotic eye–fork length=180.6 cm (CI=176–186 cm), (2) Brody growth coefficient=0.36 (CI=0.34–0.39), and (3) hypothetical age at length zero=−0.24 (CI=−0.30 to−0.18). Consequently, we believe that the von Bertalanffy growth function adequately described the individual mean growth for this species in the region.

The pelagic larva of the Midnight Snapper, Macolor macularis (Teleostei: Lutjanidae)

A larva of the Indo-West Pacific lutjanid fish Macolor macularis Fowler, the Midnight Snapper, is described from a 6.0 mm (standard length) specimen, collected in the western Pacific Ocean (2°58.1'S 158°14.3'E). Although recently settled M. macularis have been described, this is the first report of a pelagic larva. The deep-bodied larva is typical of the perciform family Lutjanidae and is characterized by 24 myomeres, laterally compressed head and body, robust, long dorsal and pelvic-fin spines with fine serrations and strong head spination. The larva of M. macularis can be distinguished from the similar larvae of Macolor niger (Forsskal) by having fin-ray counts of D X, 13 and A III, 10, the greater relative length of the second to third dorsal-fin spine and the lack of any opercular melanophores. Larvae of Macolor species are rare, juveniles are not common and adults live to 40-50 years. These facts indicate Macolor species have a life history extremely vulnerable to exploitation.

Larvae of Belonepterygion fasciolatum (Plesiopidae: Acanthoclininae)

Larvae of Belonepterygion fasciolatum (in the plesiopid subfamily Acanthoclininae) are described based on two postflexion, pelagic specimens from Espirito Santo, Vanuatu (5.7 mm SL) and Magnetic Island, Great Barrier Reef, Australia (6.3 mm SL), as well as two settled individuals (9.4 and 11.9 mm SL) from One Tree Island, Great Barrier Reef, Australia. Distinct morphological characteristics of acanthoclinins form early in development (c. 6 mm SL), including weak head spination limited to a single opercular spine, a large oblique mouth and three unbranched lateral lines. Dorsal and anal fins have many more spines than soft rays, dorsal and anal fin spines have thickened fleshy pads distally, the body is relatively slender with a deep, short caudal peduncle. Pigmentation consists of melanophores more-or-less evenly distributed across the trunk and head as well as a pale brown background (not obvious melanophores, the colour of skin and muscle) on the body that intensifies with growth. These are the first tropical acanthoclinin larvae to be reported. For reasons that are unclear, larvae of tropical and warm-temperate acanthoclinin species are rare in collections, but cool-temperate acanthoclinin larvae can be abundant.

Australoheros perdi, new species (Teleostei: Labroidei: Cichlidae) from the lacustrine region of the Doce River Valley, southeastern Brazil, with biological information

Australoheros perdi is a new species herein described from the lacustrine region of the middle Doce River basin, Doce River Valley, southeastern Brazil. The new species is distinguished from all its congeners by having only 25 vertebrae and by a combination of characters states listed below: three abdominal bars in all stages of life, a conspicuous, rounded caudalfin base spot, a conspicuous wide longitudinal stripe, head with depression in the region above the eyes, tip of pelvic fin reaching vertical through third to sixth anal fin spine base, fewer dorsal-fin spines, fewer pectoral-fin rays, fewer caudal vertebrae, fewer proximal radials on dorsal-fin base, more proximal radials on anal-fin base, more pleural ribs, a longer caudal peduncle, a deeper caudal peduncle, a wide ectopterygoid, a longer last anal-fin spine and a longer lower jaw. The available phylogenetic tree of the genus does not include the species from southeastern Brazil, thus impeding any discussion of the phylogeny of these species.

Comparison of common carp (Cyprinus carpio) age estimates derived from dorsal fin spines and pectoral fin rays

Understanding population dynamics of invasive species is critical for formulating effective management plans but dependent upon the reliability of age estimations. We compared age estimates derived from dorsal fin spines and pectoral fin rays from common carp (Cyprinus carpio), a widespread invasive species that imposes deleterious ecosystem effects. Age estimates derived from each structure were highly correlated with one another. Percentage agreement (PA) was higher (57.9%) and coefficient of variation (CV) was lower (6.6%) initially for dorsal fin spines compared to pectoral fin rays (43.5%, 9.9%, respectively), but both structures had high PA (∼95%) and low CV (0.3%) following a joint examination. Thus, both dorsal fin spines and pectoral fin rays provide similar age estimates of common carp and can be used to understand population dynamics of this species.

New species of Trachycorystes Bleeker, with comments on other species of the genus (Ostariophysi: Siluriformes: Auchenipteridae)

A new species of Trachycorystes from the rio Aripuanã, above Dardanelos and Andorinhas falls, is described. The new species is distinguished from the only other species of the genus, T. trachycorystes, by the following characteristics: jaws of equal length (vs. lower jaw prognathous in T. trachycorystes); skull roof covered by thick (vs. thin) integument; inner mental barbel very thin and short not reaching base of outer barbel (vs. extending to or beyond base of outer mental barbel); dorsal-fin spine serrated posteriorly, smooth or rough anteriorly (vs. serrated anteriorly and smooth or rough posteriorly); caudal fin shallowly forked (vs. emarginate); and gas bladder simple, without diverticula (vs. with three posterior diverticula). Comments and data on the nominal species Trachycorystes trachycorystes are provided. Trachycorystes cratensis Miranda Ribeiro, 1937, is allocated to the genus Trachelyopterus Valenciennes, 1840, and another local catfish species, Parotocinclus aripuanensis Garavello, 1988, has its type locality reassigned.

Biometric characteristics of small Atlantic bluefin tuna (Thunnus thynnus, Linnaeus, 1758) of Mediterranean Sea origin

The aim of this study was to provide a detailed description of the biometry of the juvenile Atlantic bluefin tuna (FL = 50–127 cm) of Mediterranean Sea origin. Regarding meristic features, there were 31.5 (SD = 2.1) gillrakers found on the first gill arch, 13.41 (SD = 0.494) spines in the first dorsal fin, 33.56 (SD = 2.124) soft rays in the pectoral fin, 9.86 (SD = 0.464) dorsal finlets and 8.90 (SD = 0.412) ventral finlets on the tail. Results of morphology analyses indicate that, on average, the head length contributes 28.84% (SD = 0.745); first predorsal length contributes 31.86% (SD = 0.915); preanal length contributes 61.45% (SD = 1.632); maximum body height contributes 26.90% (SD = 1.001); second dorsal fin height contributes 11.11% (SD = 1.457); pectoral fin length contributes 20.09% (SD = 1.550); and caudal fin width contributes 28.60% (SD = 1.842) of the fork length. Eye diameter comprises 13.14% (SD = 1.471), snout length 33.11% (SD = 1.332) and postorbital length 53.75% (SD = 1.354) of the head length. The significant change in body proportions, as fish grow in length, has been noted and leads to the conclusion that the juvenile bluefin body does not grow isometrically in size. Continuous decrease in the mean condition factors as body length increased, and a length–weight relationship describing negative allometric growth [ln WW (g) = −3.2965 + 2.8546 ln FL (cm)], indicate elongation of the juvenile bluefin body shape during early life, corresponding mostly to the period preceding sexual maturity.

Description of a new goatfish species, <i>Upeneus randalli</i> n. sp. (Mullidae), from the Persian Gulf, with remarks and identification keys for the genus <i>Upeneus</i>

The number of species of the goatfish genus Upeneus from the western Indian Ocean has increased considerably due to recent taxonomic efforts. Here we describe a new species, Upeneus randalli n. sp., based on body and fin coloration and 50 morphometric and meristic characters of eight specimens from the Persian Gulf, and supported by comparative data from a taxonomic review of western Indian Ocean Upeneus species and additional material from collections. Upeneus randalli is distinguished from congeners by the combination of 8 dorsal-fin spines, 14 pectoral-fin rays, 23 to 25 gill rakers, 28 to 30 lateral-line scales, pectoral-fin width 22 to 26 times in SL, and red bars on both caudal-fin lobes. We provide two identification keys: one key is for distinguishing among four species groups and the morphologically unique U. filifer , accounting for all 28 species of the genus, and the other key is for the six closely related species of the “ U. tragula group” from the western Indian Ocean, including U. randalli n. sp. Since the colour patterns fade away soon after collection and can be completely lost during preservation, morphometric and meristic characters are important for correctly identifying U. randalli n. sp. and the closely related U. margarethae . The latter species shows considerable variation in body form in different areas of the western Indian Ocean. We discuss the significance of detailed comparative studies of caudal-fin colour patterns and body-form characters for obtaining a deeper understanding of inter- and intraspecific diversity among Upeneus .

English

The study aims to identify the age and growth parameters of bullet tuna ( Auxis roche i) in Turkish waters. The fish were caught from the Turkish Mediterranean coasts between December 2008 and December 2009. A total of 186 dorsal fin spines obtained from the specimens were analyzed for ageing and growth studies. The fork length of the aged individuals ranged from 34 to 48 cm for males and from 35 to 46.5 cm for the females. Fish ages ranged 1 to 5 years old and the mean lengths by age were calculated for both sexes. Growth parameter estimates were calculated from 150 cut spine sections which provided readable growth annuli by sex. The growth parameters based on standard von bertalanffy growth function are the following: for males, L∞ (asymptotic length) is equal to 60.417 cm; k (growth coefficient) is equal to 0.159; t 0 (age at zero length) is equal to − 4.311; for females, L∞ is equal to 49.238 cm; k is equal to 0.312; t 0 is equal to −3.011; for both sexes, L∞ is equal to 57.388 cm; k is equal to 0.181and t 0 is equal to −4.155, respectively. The length and weight relationship was calculated with W equal to 0.0542 L 2.68 . Key words : Bullet tuna, Auxis rochei, age determination, growth, the Mediterranean sea.

Occurrence of the Hybodont Shark GenusMeristodonoides(Chondrichthyes; Hybodontiformes) in the Cretaceous of Kansas

Although known from Mesozoic marine, fresh and brackish water deposits from North America, Europe, Africa and Asia, hybodont shark remains have been rarely reported from the Cretaceous formations of Kansas. Remains of hybodonts reported from outside Kansas consist of small teeth, distinctive cephalic hooks, dorsal fin spines, and occasionally even complete fish. Collections of micro-vertebrate remains in Kansas since 1999 have confirmed the presence of hybodontiformes in the Kiowa (Albian), Dakota (Cenomanian) and Carlile (Turonian) formations in three different counties in the state. These specimens have been assigned to the genus Meristodonoides Underwood and Cumbaa 2010, and add an important element to the known marine faunas of the Western Interior Sea during late Early Cretaceous and early Late Cretaceous time.

Fin-Spine Selection and Section Level Influence Potential Age Estimates of Striped Marlin (Kajikia audax)

Annulus counts were compared between four sectioned levels of first dorsal-fin spines (D1–6) and anal-fin spines (A1–3) of Striped Marlin (Kajikia audax) caught in the southwest Pacific Ocean. The number of annuli observed in transverse sections varied significantly between spines of individual fish and between sections cut at varying distances along spines. Sections at levels ¼ and 1 condyle width (CW) of spines D4–6 and A3 displayed the greatest number of annuli and were not significantly different from one another. Pending validation of yearly periodicity of annulus formation, these sections could be used interchangeably in future age and growth studies on this species. The percentage of bone remodelling (vascularization) present within the central portion of fin-spine sections was negatively correlated with annulus counts. Vascularization affected the visibility of early formed annuli, which resulted in reduced counts and therefore ‘age’ estimates. Annulus counts were reduced in fin-spines D1–3, A1–2,...

Age, growth and reproduction of a common deep-water shark, shortspine spurdog (Squalus cf. mitsukurii), from Hawaiian waters

About half of the extant shark species occur only in deep waters (defined as >200 m depth), yet few published studies on sharks include these taxa. As fisheries worldwide enter deeper waters, the provision of biological data for these little-known taxa is critical to management and conservation. The shortspine spurdog, Squalus cf. mitsukurii, is an abundant shark on the insular slopes of the Hawaiian Islands. We assigned ages by counting growth bands on the enamel caps of both dorsal fin spines. Age estimates ranged from 3 to 26 years for females and from 6 to 23 years for males. Growth was modelled with multiple length-at-age models, fitted using maximum likelihood estimation and nonlinear least-squares methods. For female data, the logistic model yielded the most biologically cogent parameter estimates (L∞ = 126 cm (total length, TL) and k = 0.080 year–1). The two-parameter von Bertalanffy Growth Model yielded optimal model fit and realistic parameter estimates for males (L∞ = 72 cm (TL) and k = 0.080 year–1). Maturity ogives suggested that females and males mature at 64-cm TL (15 years) and 47-cm TL (8.5 years), respectively. Fecundity ranged from 3 to 10 embryos; mating appeared to be aseasonal. We reveal a conservative life history, common among deep-water elasmobranchs, and provide further evidence of geographic variation in reproductive and growth parameters in this nominal species.

Glyptothorax dikrongensis, a new species of catfish (Teleostei: Sisoridae) from Arunachal Pradesh, northeastern India

Glyptothorax dikrongensis, a new species of sisorid catfish from the Dikrong River in northeastern India, is described. Glyptothorax dikrongensis can be differentiated from all congeners, except G. indicus, G. rugimentum and G. obliquimaculatus, by the presence of an unculiferous patch on the posterior region of the lower lip, in between the inner mandibular-barbel bases, and unculiferous striae of the thoracic adhesive apparatus extending anteriorly onto the gular region. Glyptothorax dikrongensis is distinguished from G. indicus by the following combination of characters: equal distance between the posterior end of the pectoral-fin base and the pelvic-fin origin and between the pelvic-fin and the anal-fin origin (vs. distance between posterior end of pectoral-fin base and pelvic-fin origin greater than between pelvic-fin origin and anal-fin origin), and the pelvic-fin origin anterior to or almost at a vertical through the posterior end of the dorsal-fin base (vs. posterior to the dorsal-fin base). Glyptothorax dikrongensis is distinguished from G. rugimentum in lacking vertical bars on the body and caudal peduncle, and having a deeper caudal peduncle (8.4–9.2 vs. 6.1–7.6% SL) and a shorter dorsal-fin spine (10.1–11.1 vs. 15.2–18.6% SL). It is distinguished from G. obliquimaculatus in lacking dark, oblique blotches on the body, and in having a shorter dorsal-fin spine (10.1–11.1 vs. 13.4–16.4% SL).

Description of a new species of Microglanis from rio Barra Seca basin, southeastern Brazil (Siluriformes: Pseudopmelodidae)

Microglanis minutus, new species, is described from the rio Barra Seca basin. This species belongs to the Microglanis parahybae species complex by the following combination of characters: body light brown with fi rst and second dark brown saddles not extending ventrally below horizontal through pectoral-fin origin; light oval spot located below dorsalfin spine; third dark brown saddle not reaching anal-fin base; and caudal fin with upper lobe more developed than the inferior. The new species differs from all others M. parahybae species complex by a combination of morphological characters: pectoral and anal fins mottled or with relatively faint bands; caudal fin emarginate, trunk with dark brown saddles; caudal peduncle with faint to dark blotch irregularly shaped, caudal fin lightly mottled with narrow vertical dark brown band across central portions of lobes, dark saddle beneath adipose fin not extending ventrally to anal fin as continuous bar, continuous portion of lateral line not reaching vertical through origin of adipose fin, tip of pectoral spine as distinct bony point, a wider head, a dark blotch beneath adipose fin not extending to anal fin, distal portion of anal fin without dark band, dark saddle beneath adipose fin extending ventrally to more than the half of trunk and a light oval spot located below dorsal-fin spine.

A new specimen of the hybodont shark Palaeobates polaris with three-dimensionally preserved Meckel's cartilage from the Smithian (Early Triassic) of Spitsbergen

ABSTRACT A new, well-preserved specimen of Palaeobates polaris from the Smithian ‘fish horizon’ of Spitsbergen (Svalbard archipelago, Arctic Norway) is presented. The find is more complete than the type material of P. polaris and contains amongst others the left mandibular branch with associated dentition and labial cartilage as well as elements of the hyoid arch and a portion of the anterior dorsal fin. P. polaris shares with the other species of the genus Palaeobates the same tooth histology (orthodont teeth with pulp cavity), but the ornamentation of the teeth is different. Several studies have shown that it is problematic to deduce phylogenetic relationships among hybodontiform sharks by means of tooth histology. Moreover, orthodont teeth with a pulp cavity are probably plesiomorphic for hybodontiforms. Based on the new find of P. polaris, three other characters are proposed to be apomorphic for Palaeobates in general: (1) the lower margin of the dental groove of Meckel's cartilage runs nearer to the ventral than to the dorsal border of the mandible in P. polaris but medially to these margins of the lower jaw in the crown group hybodontoid Acrodus; (2) the number of tooth files and arrangement of the teeth of P. polaris are well-distinguishable from those of Acrodus and Asteracanthus; and (3) the dorsal fin spines of P. polaris, and also of P. angustissimus, tend to be more slender compared to those of Hybodus and Acrodus. Nonetheless, more material is needed to ascertain if these traits are useful to separate Palaeobates from other hybodontiforms.

Description of a new species of Tilapia Smith, 1840 (Teleostei: Cichlidae) from Ghana

A new species of the genus Tilapia Smith, 1840 is described from the Pra River drainage in Ghana. Tilapia pra sp. nov. is distinguished from all Tilapia species except T. sparrmanii, T. ruweti, T. guinasana, T. baloni, T. brevimanus, T. mariae, T. cabrae and T. busumana in having bicuspid posterior pharyngeal teeth on the lower pharyngeal jaw. It differs from T. baloni and T. ruweti in having more gill rakers on the first ceratobranchial (lower) gill-arch (10–12 vs. 6–9), from T. guinasana in having a higher number of upper lateral line scales (18–22 vs. 14–17) and from T. sparrmanii in a combination of a higher number of upper lateral line scales (18–22 vs. 14–19), a shorter anal fin base (15.0–18.6% vs. 18.0–23.8% of SL) and a lower number of vertical stripes (6–7 vs. 8–9). It differs from T. mariae, T. cabrae and T. brevimanus in having robust, non-spatulate outer row jaw teeth (vs. gracile spatulate teeth) and from T. busumana in having a longer last dorsal-fin spine (16.2–21.3% vs. 11.6–14.9% of SL), and a smaller lower lip length (8.0–10.7% vs. 9.6–13.9% of SL) and lower jaw length (9.9–13.6% vs. 10.5–15.2% of SL). In addition, T. pra sp. nov. differs from T. busumana in ground coloration. T. pra sp. nov. possesses a light brown to greyish dorsum and a beige to yellow ventral area vs. a bluish-purple to blackish dorsum and darker on underside of head and body of T. busumana.

Description of a new species of Tilapia Smith, 1840 (Teleostei: Cichlidae) from Ghana

A new species of the genus Tilapia Smith, 1840 is described from the Pra River drainage in Ghana. Tilapia pra sp. nov. is distinguished from all Tilapia species except T. sparrmanii, T. ruweti, T. guinasana, T. baloni, T. brevimanus, T. mariae, T. cabrae and T. busumana in having bicuspid posterior pharyngeal teeth on the lower pharyngeal jaw. It differs from T. baloni and T. ruweti in having more gill rakers on the first ceratobranchial (lower) gill-arch (10–12 vs. 6–9), from T. guinasana in having a higher number of upper lateral line scales (18–22 vs. 14–17) and from T. sparrmanii in a combination of a higher number of upper lateral line scales (18–22 vs. 14–19), a shorter anal fin base (15.0–18.6% vs. 18.0–23.8% of SL) and a lower number of vertical stripes (6–7 vs. 8–9). It differs from T. mariae, T. cabrae and T. brevimanus in having robust, non-spatulate outer row jaw teeth (vs. gracile spatulate teeth) and from T. busumana in having a longer last dorsal-fin spine (16.2–21.3% vs. 11.6–14.9% of SL), and a smaller lower lip length (8.0–10.7% vs. 9.6–13.9% of SL) and lower jaw length (9.9–13.6% vs. 10.5–15.2% of SL). In addition, T. pra sp. nov. differs from T. busumana in ground coloration. T. pra sp. nov. possesses a light brown to greyish dorsum and a beige to yellow ventral area vs. a bluish-purple to blackish dorsum and darker on underside of head and body of T. busumana.