Phylogeography and evolutionary history of the Panamic Clingfish Gobiesox adustus in the Tropical Eastern Pacific.

The Panamic Clingfish Gobiesox adustus is widely distributed in the Tropical Eastern Pacific (TEP), from the central Gulf of California, Mexico to Ecuador, including the oceanic Revillagigedo Archipelago, and Isla del Coco. This cryptobenthic species is restricted to very shallow rocky-reef habitats. Here, we used one mitochondrial and three nuclear DNA markers from 155 individuals collected across the distribution range of the species in order to evaluate if geographically structured populations exist and to elucidate its evolutionary history. Phylogenetic analyses recovered a monophyletic group, with four well-supported, allopatric subgroups. Each subgroup corresponded to one of the following well-known biogeographic regions/provinces: 1) the Revillagigedo Archipelago, 2) the Cortez + Mexican provinces (Mexico), 3) the Panamic province (from El Salvador to Ecuador), and 4) Isla del Coco. A molecular-clock analysis showed a mean date for the divergence between clade I (the Revillagigedos and Cortez + Mexican provinces) and clade II (Panamic province and Isla del Coco) in the Pliocene, at ca. 5.33 Mya. Within clade I, the segregation between the Revillagigedos and Cortez + Mexican province populations was dated at ca. 1.18 Mya, during the Pleistocene. Within clade II, the segregation between samples of Isla del Coco and the Panamic province samples was dated at ca. 0.77 Mya, during the Pleistocene. The species tree, Bayesian species delimitation tests (BPP and STACEY), the ΦST, AMOVA, and the substantial genetic distances that exist between those four subgroups, indicate that they are independent evolutionary units. These cladogenetic events seem to be related to habitat discontinuities, and oceanographic and geological processes that produce barriers to gene flow for G. adustus, effects of which are enhanced by the intrinsic ecological characteristics of this species.

The effects of soft and rough substrates on suction-based adhesion.

The northern clingfish (Gobiesox maeandricus) has a suction-based adhesive disc that can stick to incredibly rough surfaces, a challenge for stiff commercial suction cups. Both clingfish discs and bioinspired suction cups have stiff cores but flexible edges that can deform to overcome surface irregularities. Compliant surfaces are common in nature and technical settings, but performance data for fish and commercial cups are gathered from stiff surfaces. We quantified the interaction between substrate compliance, surface roughness and suction performance for the northern clingfish, commercial suction cups and three biomimetic suction cups with disc rims of varying compliance. We found that all cups stick better on stiffer substrates and worse on more compliant ones, as indicated by peak stress values. On compliant substrates, surface roughness had little effect on adhesion, even for commercial cups that normally fail on hard, rough surfaces. We propose that suction performance on compliant substrates can be explained in part by effective elastic modulus, the combined elastic modulus from a cup-substrate interaction. Of all the tested cups, the biomimetic cups performed the best on compliant surfaces, highlighting their potential to be used in medical and marine geotechnical fields. Lastly, we discuss the overmolding technique used to generate the bioinspired cups and how it is an important tool for studying biology.

A multi-locus approach to elucidating the evolutionary history of the clingfish Tomicodon petersii (Gobiesocidae) in the Tropical Eastern Pacific.

Marine species that are widely distributed in the Tropical Eastern Pacific (TEP) has served as a model for studying biogeographic patterns resulting from the effects of intraregional habitat discontinuities and oceanographic processes on the diversification and evolution of cryptobenthic reef fishes. Tomicodon petersii, a clingfish (Gobiesocidae) endemic to the TEP, is found on very shallow rocky reefs from central Mexico to northern Peru, and in the Cocos and Galapagos islands. We evaluated the effect of likely biogeographic barriers in different parts of the TEP on the diversification process of this species. We used one mitochondrial and three nuclear DNA markers from 112 individuals collected across the distribution range of T. petersii. Our phylogenetic results showed the samples constituted a monophyletic group, with three well-supported, allopatric subgroups: in the Mexican province, the Panamic province (from El Salvador to Ecuador), and the Galapagos Islands. The split between the Mexican and more southerly clades was estimated to occur at the end of the Miocene ca. 5.74 Mya, and the subsequent cladogenetic event separating the Galapagos population from the Panamic population at the junction of the Pliocene and Pleistocene, ca. 2.85 Mya. The species tree, Bayesian species delimitation tests (BPP), STACEY, and substantial genetic distances separating these three populations indicate that these three independent evolutionary units likely include two unnamed species. The cladogenetic events that promoted the formation of those genetically differentiated groups are consistent with disruptive effects on gene flow of habitat discontinuities and oceanographic processes along the mainland shoreline in the TEP and of ocean-island isolation, in conjunction with the species intrinsic life-history characteristics.

First documentation of the otoliths of the species of Gouania (Teleostei: Gobiesocidae) in the Mediterranean Sea.

Otolith morphology is a widely accepted tool for species identification in teleost fish, but whether this holds true for very small species remains to be explored. Here, the saccular otoliths of the cryptobenthic Mediterranean clingfish Gouania (Gobiesocidae) are described for the first time. The new data, although preliminary, indicate that otolith morphology and morphometry support the recognition of the recently differentiated five species of Gouania in the Mediterranean Sea. Furthermore, otoliths of phylogenetically closely related Gouania species resemble each other more than do those of the more distantly related species.

Unravelling the taxonomy of an interstitial fish radiation: Three new species of Gouania (Teleostei: Gobiesocidae) from the Mediterranean Sea and redescriptions of G. willdenowi and G. pigra.

The clingfish (Gobiesocidae) genus Gouania Nardo, 1833 is endemic to the Mediterranean Sea and inhabits, unlike any other vertebrate species in Europe, the harsh intertidal environment of gravel beaches. Following up on a previous phylogenetic study, we revise the diversity and taxonomy of this genus by analysing a comprehensive set of morphological (meristics, morphometrics, microcomputed tomography imaging), geographical and genetic (DNA-barcoding) data. We provide descriptions of three new species, G. adriatica sp. nov., G. orientalis sp. nov. and G. hofrichteri sp. nov., as well as redescriptions of G. willdenowi (Risso, 1810) and G. pigra (Nardo, 1827) and assign neotypes for the latter two species. In addition to elucidating the complex taxonomic situation of Gouania, we discuss the potential of this enigmatic clingfish genus for further ecological, evolutionary and biodiversity studies that might unravel even more diversity in this unique Mediterranean fish radiation.

Larval parasitic copepods affect early life history traits of a temperate clingfish.

Larval copepods are frequent parasites that infest fish larvae along the Chilean coast. Because these parasites develop on fish during the early development, when their bodies are fragile and in a recent stage, they can affect the fishes' early life history traits (ELHT). The goal of this study was to determine the effect of parasitic copepods on the ELHT of the larvae of the clingfish Gobiesox marmoratus (Teleostei: Gobiesocidae) using otolith microstructure analysis. Ichthyoplankton samples were collected during austral winter (July and August 2012), in the inner shelf waters off Valparaiso Bay, central Chile. A total of 95 non-parasitized larvae (NPL) and 95 parasitized (PL) with copepods were randomly selected for subsequent analyses. Parasitized larvae of G. marmoratus were larger than NPL. The right otolith tended to be larger than the left otolith in the fish larvae, but with a higher asymmetry in PL. The PL showed larger otoliths-at-size than the NPL, particularly in smaller larvae (< 8 mm of standard length, SL). Nonetheless, parasitized larvae larger than 8 mm SL showed the opposite trend that is smaller-at-size otoliths than NPL. The Gompertz models indicated that the asymptotic length of NPL doubled the length of PL; this suggests that parasitic copepods affect the maximum size attained by the PL. In conclusion, parasitic copepods negatively affect the ELHT of G. marmoratus larvae and the greater asymmetry can be attributed to parasitism.

Independent evolutionary lineage of the clingfish Gobiesox adustus (Gobiesocidae) from Isla del Coco, Costa Rica

Introduction: Isla del Coco is an important protected area for marine fauna in the Eastern Tropical Pacific. In this area, the species that inhabit the intertidal zone have been subject to few studies. One of the species inhabiting these areas is the clingfish Gobiesox adustus (Gobiesocidae). Objective: To analyze for the first time the mitochondrial gene cytochrome oxidase subunit 1 (cox1) of G. adustus' population from Isla del Coco and compare it with those of continental coast of Costa Rica and Ecuador. Methods: We constructed a haplotype network for these samples. Genetic diversity, distance and structure were calculated by several software. The historical demography of Isla del Coco samples was assessed with the method Bayesian skyline plot as implemented in BEAST2. Results: The samples segregate into three haplogroups: one consisting of the Isla del Coco samples, a second consisting of a subset of the Ecuador samples, and a third consisting of Costa Rica and the remaining Ecuador samples. The genetic distances between the three haplogroups range between 1.6% and 2.1% (uncorrected p-distance), and pairwise ΦST and AMOVA results between the three haplogroups show high and significant values. Conclusions: The Isla del Coco haplogroup showed a Pleistocene population growth, which agrees with demographic patterns found in other marine organisms. The history of isolation of the G. adustus population from Isla del Coco demonstrates the evolutionary independence of this population.

Introducción: Isla del Coco es un área protegida importante para la fauna marina en el Pacifico Oriental Tropical. En esta área, las especies que habitan la zona intermareal han sido objeto de pocos estudios. Una de las especies que habitan en estas áreas es el clingfish Gobiesox adustus (Gobiesocidae). Objetivo: Analizar por primera vez el gen mitocondrial citocromo oxidasa sub unidad 1 (cox1) de poblaciones de Isla del Coco y compararlo con las de la zona continental de Costa Rica y Ecuador. Métodos: Se construyó una red de haplotipos. La diversidad, la distancia y la estructura genética fueron calculadas por diversos programas. La demografía histórica de las muestras de Isla del Coco fue evaluada con el método Bayesian skyline plot implementado en BEAST2. Resultados: Las muestras se agruparon en tres haplogrupos: en un haplogrupo se incluyó a los individuos de Isla del Coco, otro haplogrupo integró las muestras de Ecuador y un tercer grupo incluyó las muestras restantes de Costa Rica y Ecuador. Las distancias genéticas entre los tres haplogrupos oscilan entre 1.6% y 2.1% (p-distancia, no corregida), las distancias ΦST y los resultados de AMOVA entre los tres haplogrupos muestran valores altos y significativos. Conclusiones: El haplogrupo de Isla del Coco mostró un crecimiento poblacional datado en el Pleistoceno, coincidiendo con la demografía poblacional encontrada en otros organismos marinos. La historia de aislamiento de la población de G. adustus de Isla del Coco demostró la independencia evolutiva de esta población.

Validity of Lepadichthys misakius (Tanaka 1908) and redescription of Lepadichthys frenatus Waite 1904 (Gobiesocidae: Diademichthyinae).

The clingfishes Lepadichthys frenatus Waite 1904 and Lepadichthys misakius (Tanaka 1908) are both redescribed as valid species, although the latter has previously been regarded as a junior synonym of the former. Lepadichthys frenatus and L. misakius are easily distinguished from their congeners by the following combination of characters: 14-19 dorsal-fin rays; 12-14 anal-fin rays; disc size moderate, its length 15.1-20.8 % of standard length (SL); and dorsal, anal and caudal fins connected by membranes. Lepadichthys misakius can be distinguished from L. frenatus by having 25-28 (modally 26 or 27) pectoral-fin rays [vs. 27-29 (29) in L. frenatus]; the upper end of the gill membrane level with the 5th to 8th (usually 6th) pectoral-fin ray base in lateral view [vs. 6th to 8th (7th)]; the lower 8th to 11th (9th) pectoral-fin ray base attached to the disc base by membrane [vs. 10th to 12th (11th)]; 7-11 (9) gill rakers on each arch [vs. 11-14 (12)]; anterior, posterior and least interorbital widths 9.0-11.8 (mean 10.7), 13.4-16.9 (15.3) and 3.2-8.8 (6.2) % SL, respectively [vs. 8.4-10.2 (9.4), 12.1-14.7 (13.5) and 2.8-5.9 (4.8) % SL, respectively]; and NC2 (nasal canal pore) usually located between the anterior and posterior margins of the posterior nostril (vs. usually located before the posterior nostril anterior margin). Morphological changes with growth of the two species are described in detail and a lectotype designated for L. frenatus.

Parasitic copepods affect morphospace and diet of larvae of a temperate reef fish.

The effects of ectoparasites on larvae of the clingfish Gobiesox marmoratus were evaluated at the dietary and morphometric levels. The larvae and ectoparasites were collected by nearshore plankton samplings during October, November and December 2013 off El Quisco Bay, central Chile. The standardized abundance of total larvae and those ectoparasitized larvae (PL) was positively related and high parasite prevalence was found throughout the sampling period (up to 38%). Geometric morphometrics analyses indicate main changes in the shape through early ontogeny and subtle but significant variations between PL and non-parasitized larvae (NPL). Prey composition varied between PL and NPL; small size (<6 mm standard length, LS ) parasitized larval G. marmoratus ate mostly gastropod larvae, whereas small non-parasitized specimens ate mainly cirripede nauplii. All larger (>8 mm LS ), pre-settlement stages parasitized by Trifur and, or Caligus copepods had content in their gut, suggesting that ectoparasites did not diminish prey capture in host with larger size. Morphometric and dietary changes occurring during larval development were decoupled, both for PL and NPL. The maintenance of a slender, more hydrodynamic body through pelagic development and the ingestion of less-mobile prey in PL suggests non-lethal effects of ectoparasitism on rocky-reef fish larvae.