Dorsal-fin Base Research Articles

The ornate rubbernose pleco (Siluriformes, Loricariidae, Chaetostoma), a new species from the Ucayali River Basin, Peru

A new species in the rubbernose catfish genus Chaetostoma is described from the Aguaytia, Pisqui and Palcazu Rivers, which drain the Pampa de Sacramento Region in the Ucayali River drainage of central Peru. The new species is distinguished from congeners, except C. anomalum, C. branickii, C. dorsale, C. leucomelas, C. microps, C. nudirostre, C. palmeri and C. thomsoni by having distinct, white, variably-shaped spots or vermiculations ½–2× nostril diameter on dark grey to black background on the head (vs. spots absent or black on light-coloured background). The new species is distinguished from C. anomalum, C. branickii, C. dorsale, C. microps, C. nudirostre and C. thomsoni by having highly variable, distinct white spots, vermiculations or bands ½–10× nostril diameter on the body, from C. leucomelas by having dorsal and caudal fin indistinctly and variably-patterned with zero to four bands (vs. dorsal and caudal fin consistently having five or more uniform bands) and from C. palmeri by having two predorsal plates (vs. three), supraoccipital excrescence present (vs. absent) and pelvic-fin insertion slightly posterior to dorsal-fin insertion (vs. pelvic-fin insertion at middle of dorsal-fin base). Species delimitation analyses of the COI and Cytb genes further support the recognition of this new species.

Gaidropsarus mauritanicus (Gadiformes, Gaidropsaridae) a new three-bearded rockling from a deep-water coral ecosystem with a genetically verified biogeographical distribution of the genus and notes to its ecology and behavior.

Gaidropsarus mauritanicus sp. nov. is described from one specimen collected using a grab sample from the Tanoûdêrt Canyon (ca. 20° N) off Mauritania at a depth of 595 m. The new species was further observed during eight remotely operated vehicle (ROV) dives along the Mauritanian slope southwards down to the Tiguent Coral Mound Complex (~17° N) in a bathymetric range between 613 and 416 m. It can be distinguished from congeners by a combination of characteristics, including large eyes (38.1% head length [HL]), large head (25.8% standard length [SL]), elongated pelvic fins (35.7% SL), low number of vertebrae (44), and coloration (pinkish with a dorsal darker brownish hue and bright blotches along the dorsal-fin base). A species-delimitation analysis performed with available cytochrome c oxidase subunit 1 (COI) sequences affiliated to the genus Gaidropsarus additionally supported the validity of the new species. Video analyses showed a deep-water coral-associated and protection-seeking behavior, which may explain why the species has remained undescribed until now. Additional ROV footage from separate deep-water coral sites in the North Atlantic and Mediterranean Sea further highlights the ecological behavior and hidden diversity of bathyal three-bearded rocklings. Here, we additionally discuss the biogeographical distribution of all genetically verified Gaidropsarus spp. in combination with genetic data and morphological characters. G. mauritanicus sp. nov. is closely related to a species from Tasmania (43° S), a geographical point furthest among the studied samples, which may hint to an important influence of (paleo-) oceanography on the distributions of Gaidropsarus species.

Cryptic diversity of mudskipper genus Boleophthalmus (Gobiiformes: Oxudercidae) from the north coast of East Java, Indonesia

Abstract. Shabrina FN, Wibowo K, Arisuryanti T. 2024. Cryptic diversity of mudskipper genus Boleophthalmus (Gobiiformes: Oxudercidae) from the north coast of East Java, Indonesia. Biodiversitas 25: 412-420. Mudskippers of genus Boleophthalmus are commonly found in the Indo-Pacific area. This study aimed to examine the diversity of the genus Boleophthalmus from the north coast of East Java Province, Indonesia by employing the mitochondrial COI gene for DNA barcoding and morphological characterization. Ten individuals of this genus were analyzed. According to the similarity percentage evaluated using Basic Local Alignment Search Tool (BLAST) and Barcode of Life Data (BOLD) identification engines, all the individuals were identified as Boleophthalmus boddarti Pallas, 1770. However, phylogenetic analysis revealed that three of the individuals were identified as B. boddarti, and seven were identified as Boleophthalmus pectinirostris Linnaeus, 1758 originated from East Java, which is separated from the clade of B. pectinirostris from East Asia population. These results were supported by the genetic distance of 8.93%-13.81% and the morphological characteristics determined by principal component analysis (PCA). PCA revealed that B. boddarti and B. pectinirostris are primarily distinguished by the number of (longitudinal) scales (69-75 versus 113-123), length of second dorsal fin base (42.3-43.9 versus 38.1-49.0), and caudal fin length (18.1-22.4 versus 17.5-21.6). Further detailed studies are needed to clarify whether the seven individuals belonging to B. pectinirostris complex should be regarded as a new species or a new subspecies.

Evaluation of body performance of tilapia (Oreochromis niloticus)

Efforts to improve the quality of fish seeds can be done by using quality broodstock which includes having the right body shape. This study aimed to evaluate the body performance of male and female Sultana strain tilapia (Oreochromis niloticus) which were to be used in hatchery activities. A total of 35 male and 35 female broodstock were evaluated for body size using a ruler. Measurements of 12 morphological characters: total length (TL), standard length (SL), head length (HL), pre-dorsal length (PrDL), pre-pectoral length (PrPL), pre-pelvic length (PrPeL), pre-pelvic length (PrPeL), anal length (PrAL), anal fin length (AFL), dorsal fin base length (DFL), body height (BH), caudal peduncle height (CPH), and dorsal anal length distance (DAL). The ratio between the character values of SL, HL, PrDL, PrPL, PrPeL, PrAL, AFL, DFL, BH, CPH, DAL, and TL were analyzed descriptively. The results of the analysis showed that the average value of the ratio of length characters (SL/TL, HL/TL, DF, and AFL/TL), height character (BH/TL), and diagonal character (DAL/TL) were higher in the male broodstock than in the female broodstock. On the other hand, other length characters (PrDL/TL, PrPL/TL, PrPeL/TL, PrAL/TL) and height characters (CPH/TL) had the same values in the male and female broodstock. In general, the size of the male broodstock used tended to be larger than the female broodstock.

Short-beaked common dolphin (Delphinus delphis) total length estimation using laser photogrammetry off the southwest coast of Ireland

Measurements from 106 stranded short‐beaked common dolphins along the Irish coast were taken between March 2017 and March 2023. Data were collected from the Irish Necropsy Project and Irish Cetacean Stranding Scheme. Total length measurements were gathered from 103 individuals where the tail flukes were still attached. These ranged between 96–238cm, mean 185.7cm, SD 31.43cm. Males (n = 58) ranged from 96–238cm, averaging 189cm, SD 34.64cm. Females (n = 40) measured 117.5–231cm, averaging 181cm, SD 25.80cm. Scaled dorsal fin photos were taken from stranded dolphins to formulate an equation to estimate the total length of live dolphins. The total lengths of 29 live common dolphins were estimated off the southwest coast of Ireland using laser photogrammetry and dorsal fin dimensions between March 2018–April 2019. Total lengths for three stranded dolphins with amputated tail flukes were also estimated. Dorsal fin base lengths were the most accurate predictor of total length R2 = 0.78. Total length estimates ranged between 143.77–242.25cm, averaging 194.78cm, SD 20.05cm. The adoption of laser photogrammetry as a measurement tool warrants further exploration as a means to reduce potential disruption from aerial systems and enhance the utility of behavioural and photo‐ID images. This study describes a non‐invasive technique with a range of possible applications for understanding pod size structure and seasonality due to this species’ approachable behaviour and inquisitive nature.

Glyptothorax siangensis, a New Species of Catfish (Sisoridae) from the Brahmaputra River Drainage, Arunachal Pradesh, India

Glyptothorax siangensis, new species, is described from the Siang River in Arunachal Pradesh, India. The new species can be distinguished from its congeners from the Ganga-Brahmaputra drainage and its neighbouring drainages by the following combination of characters: a thoracic adhesive apparatus with skin ridges present over almost the entire apparatus, including the sword-shaped median depression, which is narrowly opened posteriorly; an elongated dorsal spine distinctly greater than body depth at its origin, posteriorly serrated with 11–17 serrae; nuchal plate saddle-like with W-shaped extensions; shallow body depth; long nasal barbel, almost reaching the anterior margin of eye; a complete longitudinal black band across the distal half of dorsal-fin; and tips of neural spines extension visible as a row of ridges between dorsal-fin base and origin of procurrent rays of caudal-fin. This is the nineteenth species of Glyptothorax known from the Ganga-Brahmaputra drainage.

HABITUS COMPARASION, GROWTH PATTERN AND CONDITION OF CONVICT CICHLID (Amatitlania nigrofasciata GUNTHER, 1986) IN THREE LAKES ECOSYSTEMS IN BALI

Convict cichlid (Amatitlania nigrofasciata) belonging to the Cichilidae family has inhabited in Tamblingan Lake, Buyan Lake and Beratan Lake. The existence of this fish species is suspected due to accidental introduction activities. The aims of this study ware to reveal differences in habitus, growth patterns and conditions of convict cichlid in the three lakes on the island of Bali. Samples of convict cichlid were taken by bubu traps, that placed in the littoral zone of the lake. 19 morphometric characters were measured using a digital caliper, then each fish sample was weighed. The growth pattern and condition of convict cichlid are known by using the equation of the length-weight relationship and the condition factor. There were 888 individuals of convict cichlid caught during this study, consisted of 385 fish from Tamblingan Lake, 371 fish from Buyan Lake, and 132 fish from Beratan Lake. Moreover, 108 fish consisting of 36 fish from each lake were further analyzed by measuring 17 additional morphometric characters. The results of measuring the morphometric characters of the observed convict cichlid had different variations for the characters of head length and predorsal length. There was diversity for several morphometric characters of the convict cichlid in Buyan Lake which are a collaboration of characters between the convict cichlid in Tamblingan Lake and in Beratan Lake, namely on the basic characteristics of the anal fin, dorsal fin base, body length, postorbital length, and tail height. The growth pattern of this species in the three lakes was negative allometric and the condition of this fish species was in good condition with the average value of condition factor ranging from 0.95–1.05.

A new species of torrent catfish, Liobagrusgeumgangensis (Teleostei, Siluriformes, Amblycipitidae), from Korea.

In a recent survey of populations of the Korean torrent catfish Liobagrus, a distinctive species was discovered from the Geum River and its tributaries flowing into the western coast of Korea, and here described as a new species, L.geumgangensissp. nov. It is distinguishable from other congeners by a combination of the following characters: I, 8 pectoral fin-rays; 52-56 caudal-fin rays; a relatively short occiput to dorsal-fin origin distance (6.9-9.8% SL); a short pelvic-fin insertion to anal-fin origin distance (11.9-17.3% SL); a long dorsal-fin base (10.6-13.5% SL); 8-9 gill rakers; 5-8 serrations on the pectoral fin; the body and fins are dark yellow, the margins of the dorsal, anal, and caudal fins are dark brown, but the outermost rim is faintly yellow. Analysis of the cytb gene also confirmed that L.geumgangensis is a monophyletic lineage distinct from other congeners.

Morphological features and length-weight relationships of <i>Sarotherodon galilaeus</i> from Oyan Lake, Southwestern Nigeria

Oyan Lake is one of the major sources of fish protein supply to many parts of Southwestern Nigeria. Investigation was conducted on the biometrics, length-weight relationship and condition factor of Sarotherodon galilaeus in Oyan Lake. Fish samples were obtained from fishermen using gillnets, hook and line, cast nets and traps. One hundred and thirty samples of S. galilaeus were collected for the study, comprising eighty-three (83) males and forty-seven (47) females with sex ratio of 1:0.57 (male:female). The mean total lengths for combined sexes, males and females were 15.5±0.15, 15.7±0.19 and 15.3±0.26cm while the mean total weights were 81.0±2.78, 83.8±0.56 and 75.9±4.70g, respectively. No significant difference (p>0.05) was observed between mean total lengths of males and females. However, mean weight of males was significantly higher (p<0.05) than females. Mean eye diameter (EYD) and length of caudal peduncle (LCP) were significantly higher (p<0.05) in females. The range of number of pectoral rays (PRA) (12-14), number of pelvic rays (PVR) (4-5), number of anal spines (ANS) (2-3) and number of pelvic spines (PEVS) (1, no range) were the same in both sexes. There was strong correlation between weight/total length (r=0.93) and total length/length of dorsal fin base (r=0.72), while there was no correlation between body depth/head length. The mean condition factor (k) were 2.07±0.02 (combined sexes), 2.08±0.03 (males) and 2.03±0.03 (females). Growth was isometric for both sexes: males, 3.24 and females, 3.06. The results suggested that Oyan Lake is suitable for S. galilaeus.

Jenis Kelamin Ikan Nilem (Osteochilus hasseltii) Berdasarkan Truss Morfometrik di Sungai Banjaran, Kabupaten Banyumas

Osteochilus hasseltii belongs to the Cyprinidae family and is a native species in the Banjaran River. Populations of O. hasseltii that are continuously exploited can cause the population to decrease and cause an imbalance between males and females. This study aims to determine differences in morphology and sex of O. hasseltii. The research method uses a survey method and purposive random sampling at two stations, namely Kober Village and Kedung Wuluh Village. A total of 16 individuals of O. hasseltii (8 males and 8 females) were caught using fishing rods. About 24 morphometric truss distances were measured using Image-J software. Normality and homogeneity tests need to be done before conducting the t-test. Data analysis used the t-test with the help of Minitab software. The morphological characters of O. hasseltii seen from the dendogram can be seen that each truss distance measured on male O. hasseltii has a similarity of above 90% so that the morphological differences and the similarities are small. Whereas in female O.hasseltii, the similarity exceeds 75% at each truss distance so that the distinguishing character of female O.hasseltii can be seen from the truss distance which has a small similarity, namely Anal-fin base length (Q) of 76.14% and Upper-jaw length (D) of 86.3%. The results of the t-test showed 8 morphometric truss distances that could be used as a distinguishing feature of male and female O. hasseltii is Snout-supraoccipital distance, Snout length, Snout-pectoral distance, Snout-gill distance, Postorbital head length, Gill-pelvic distance, Dorsal-pelvic distance, and Dorsal fin base length. Therefore, the morphometric truss can be used to differentiate the sex of O. hasseltii.

Adult morphology and redescription of Lestidiopsindopacificus (Ege, 1953), with comments on the features of related species (Teleostei, Aulopiformes, Paralepididae).

Two specimens representing the first known adults of Lestidiopsindopacificus (Ege, 1953) are reported and described from Taiwan, and the validity and generic assignment of this species are confirmed. The origin of the pelvic fin directly below the dorsal-fin base shows that L.indopacificus belongs to the L.mirabilis species complex. It can be separated from its congeners by the position of the nostrils above the posterior end of the maxilla, the light body color with unevenly distributed melanophores in adults, and a distinct combination of meristic values and other morphological characteristics. New geographic records are reported for the two other current members of this species complex, L.mirabilis (Ege, 1933) and L.extremus (Ege, 1953). The diagnostic features that separate these three very similar species are discussed.

A new species of Cambeva (Siluriformes: Trichomycteridae) from an area of high anthropogenic impacts in the headwaters of Rio Iguaçu, Southern Brazil

Cambeva piraquara, sp. nov., a restricted-range and rare species last collected from over 20 years ago, is described from the Rio Piraquara, upper Rio Iguaçu basin, Rio Paraná (La Plata) system. The new species is distinguished from all congeners by having two conspicuous dark-brown longitudinal stripes on the inner skin layer of body over a plain yellowish background: a conspicuous wide and well-defined dark-brown longitudinal mid-lateral stripe extending from the opercular patch of odontodes to the first third of caudal-fin rays and a dorso-sagittal stripe comprising large and coalescent rounded blotches extending from occipital to the dorsal-fin base. We provide an illustrated osteological description and assess the conservation status of the new species, which faces several environmental impacts in the upper Rio Iguaçu basin.

Comparative Morphometric Analysis of the Azov Anchovy (<i>Engraulis encrasicolus maeoticus</i> Pusanov, 1926) and the Black Sea Anchovy (<i>E. encrasicolus ponticus</i> Alexandrov, 1927) Based on the Body Shape

A comparative analysis of body shape was performed on 841 specimens of anchovy collected from 21 catches from the Sea of Azov, the Kerch Strait, and the Black Sea off the Crimea and the northeastern coast of the Caucasus in 2006–2007 and 2016–2017. Twenty-three measurements between 12 homologous landmarks of the body of each specimen according to the “Truss Network System” served as characters for analysis. Fifteen characters showed significant differences between the Azov anchovy Engraulis encrasicolus maeoticus Pusa-nov, 1926 and the Black Sea anchovy E. encrasicolus ponticus Alexandrov, 1927. Specimens of the Azov subspecies had larger relative values for all characters of the head region, the height of the trunk region, the re-lative length of the bases of the dorsal and anal fins, and the height of the base of the caudal fin. Specimens of the Black Sea subspecies had a greater postdorsal length and a larger distance from the end of the dorsal fin base to the end of the anal fin base. A multivariate principal component analysis revealed the highest variabi-lity of characters of body height, which depends on the fatness of the fish and demonstrates seasonal dyna-mics. The second group combines characters that do not have a seasonal component (dimensions of the head region, lengths of the bases of the dorsal and anal fins, and postdorsal distance) and can be used for distinguishing the shape of the Azov and Black Sea anchovies. The success rate of classification of Azov and Black Sea anchovies according to the body shape measurement characters using the discriminant analysis was 83.4%.

A new species of Loricaria (Loricariidae: Loricariinae) from the upper Amazon River basin, Colombia

Abstract A new species of Loricaria is described from the upper Amazon River basin, Colombia. The new species is distinguished from its congeners primarily by having the dorsal portion of head with uniform black or dark brown coloration extending to three or four plates posterior to dorsal fin base, or with two longitudinal bands from tip of the snout to origin of dorsal fin; abdominal plates tightly joined and completely covering the median abdominal space and pectoral girdle; and pectoral and dorsal fins totally black or dark brown, without bands, spots, or blotches. The new species is further distinguished by plate counts, and body measurements. An analysis of genetic distances using the cytochrome oxidase c subunit 1 marker of the mitochondrial genome showed a clear differentiation between the new species and Loricaria cataphracta (5.8–7.6%), L. nickeriensis (5.7–6.1%), and L. simillima (2.7–7.0%). Species delimitation analyses were carried out, which further supported the new species as a divergent lineage within the genus. Fish species diversity of the upper Amazon River basin and taxonomic issues related to L. simillima are included as part of the discussion.

A mismatch between morphological and molecular data in lineages of Enteromius (Cypriniformes: Cyprinidae) from the Lowa basin (East Democratic Republic of the Congo: DRC) with the description of a new species

Six populations of morphologically similar Enteromius specimens with a weakly thickened and serrated dorsal spine from the Lowa basin were investigated using an integrative approach including qualitative characters, multivariate morphometrics, and DNA analyses (mtCOI and ncMyh6). These populations are nested within the COI lineage of the E. miolepis species-complex, but differ from all known Enteromius species from the Congo basin, by their dorsal spine morphology, colour pattern, and scale pattern at the dorsal-fin base. In addition, they represent two distinct COI lineages (genetic distance: 5.6–6.1%), with four populations forming a single well-supported lineage described as a new species, Enteromius radari sp. nov. The two remaining populations are sister to the E. miolepis lineage from the Itimbiri. The incongruent position of these two populations in the COI tree may point to past introgressive hybridization events, incomplete lineage sorting, or recent speciation. Analysis of a ncDNA marker (Myh6) did not provide more information due to its slow mutation rate. These results are indicative for introgression or mtDNA capture and provide the first report for such an evolutionary event in Enteromius from the Congo basin. http://zoobank.org/urn:lsid:zoobank.org:pub:D2263E5B-D403-404C-B66C-852215ED100C

A new species of Sturisoma Swainson, 1838 (Loricariidae: Loricariinae), from the Madeira River basin, with a discussion of historical biogeography of western Amazonas and Paraguay River basins.

A new loricariin species of Sturisoma is described from the Cautário, Guaporé, Mamoré, Machado and Soteiro rivers, Madeira River basin, in Bolivia and Brazil. The new species is distinguished from its congeners by the presence of a middorsal longitudinal, thin dark brown stripe on the caudal peduncle, extending from two or three plates posterior to the dorsal-fin base, reaching the origin of the caudal fin, or one or two plates anterior to the origin of the caudal fin; small squarish anteriormost abdominal plates; and a middorsal longitudinal dark-brown stripe from first predorsal plate to near the dorsal-fin origin. Furthermore, the new species is diagnosed from congeners by plate morphology, counts on the median series, coalescent plates and ventrolateral thoracic plates, in addition to measurements related to body and head structures. An analysis of genetic distances using cytochrome C oxidase subunit 1 gene marker of the mitochondrial genome between the new species and several congeners is presented, in addition to a likelihood analysis to illustrate the position of the new taxon within Sturisoma. An identification key for species of the genus currently recorded at the upper Amazonas River basin is provided.

First adult record of the barracudina Lestidiops extremus (Ege, 1953), based on a specimen from the Philippines (Teleostei: Paralepididae)

The adult morphology of Lestidiops extremus (Ege, 1953), a barracudina originally described based on postlarvae from Indonesia, has long been a mystery. An adult specimen collected from off the Philippines represents the first confirmed record of an adult of this species. Observations on the holotype and a detailed description of the adult are provided. The location of the pelvic-fin origin in the adult, clearly behind the posterior end of the dorsal-fin base, is unique within this genus and the entire subfamily Lestidiinae. The present find extends the known range of the species from northwestern New Guinea, Indonesia, to seas east of the Philippines.

Analysis of Morphometric and Gene of Bombay-duck (Harpodon nehereus) in Tarakan Waters

Bombay-duck (Harpadon nehereus) was one of economically valuable fish at Tarakan waters. This fish was caught by trawl and jermal. In this study, samples were taken from two fishing area, that were Juata and Amal. 20 characters morphometric was measure carried out on 100 samples from each area. Molecular identification and phylogeny of this species used Cytochrome C Oxidase Subunit I (COI) gene. The phylogenetic tree was constructed used MEGA 5.0 program. There were 5 significantly different characters between sample from Juata and Amal, that were body weight (P = 0.00), distance between pelvic and anal fins (P = 0.001), caudal fin height (P = 0.009), pelvic fin base length (P = 0.015), and dorsal fin base length (P = 0.031). Molecular identification showed that samples were the same species. The genetic distance between samples from Juata and Amal was 0 (zero).

Distribution and morphobiological characteristics of the chinese false gudgeon Abbottina rivularis (Basilewsky, 1855) in the Ile river basin (Balkhash basin, Republic оf Kazakhstan

As a result of intentional and accidental introduction of alien fish species in most reservoirs of Southern and South–Eastern Kazakhstan, there was a significant change in the composition of the ichthyofauna. Alien low-value fish species have become numerous in many small reservoirs of the basin. In 2019-2021, the distribution of the river abbottina Abbottina rivularis (Basilewsky, 1855) was studied in several small reservoirs of the Ile River basin. The purpose of the research: to study the morphobiological characteristics of two populations of river abbotina in the basin of the Ile and give an assessment of their condition. Biological and morphological analyses of samples were carried out according to the most common scheme. The variability of 33 plastic and 7 counting features of two samples was studied. An indicator of fluctuating asymmetry was used to assess the conditions of existence. As a result of morphometric measurements of fish, out of 20 indicators of plastic measurements, significant changes were found in 9 (45%) plastic signs compared to the mother reservoir. In fish, the antedorsal distance (aD) increased by 6%, the length of the dorsal fin base (lD) by 25%, the height of the dorsal fin (hD) by 6.3%, the length of the anal fin (lA) by 51.4%, the length of the pectoral fins (lP) by 10.8%, the length of the ventral fin (lV) increased by 18%. The length of the head (c) increased by 11.2%, the highest body height (H) decreased by 5.3%, the lowest body height (h) increased by 26.4%. Despite significant violations of homeostasis in development, fish in ponds reach large sizes.Тhe Bakanas irrigation irrigation system can be a supplier of juvenile river abbotina for the Ile River. The size and weight indicators of the studied populations of the river abbotina from two samples showed lower values when compared with similar data from the Issyk River (1975) and the mother reservoir of the Amur River (1956). Key words: acclimatization, alien species, river abbotina, Balkhash – Ili basin, small reservoirs, morphometric characteristics, fluctuating asymmetry.

Descriptors of a New Morphotype of One Stripe Spiny Eel, Macrognathus aral(Bloch & Schneider, 1801), Family: Mastacembelidae

The new ocelli variant of Macrognathus aral (Bloch & Schneider, 1801) under Mastacembelidae family was recorded from Gangetic Wetlands of North 24 Parganas, West Bengal, India. The new variant can be considered as a potential ornamental fish apart from high valued food fish, due its more number of ocelli in both dorsal and anal fin base of the body surface, which comparable to other commercially important M. aral.