Phylogenetic relationships and divergence dates of softshell turtles (Testudines: Trionychidae) inferred from complete mitochondrial genomes

The softshell turtles (Trionychidae) are one of the most widely distributed reptile groups in the world, and fossils have been found on all continents except Antarctica. The phylogenetic relationships among members of this group have been previously studied; however, disagreements regarding its taxonomy, its phylogeography and divergence times are still poorly understood as well. Here, we present a comprehensive mitogenomic study of softshell turtles. We sequenced the complete mitochondrial genomes of 10 softshell turtles, in addition to the GenBank sequence of Dogania subplana, Lissemys punctata, Trionyx triunguis, which cover all extant genera within Trionychidae except for Cyclanorbis and Cycloderma. These data were combined with other mitogenomes of turtles for phylogenetic analyses. Divergence time calibration and ancestral reconstruction were calculated using BEAST and RASP software, respectively. Our phylogenetic analyses indicate that Trionychidae is the sister taxon of Carettochelyidae, and support the monophyly of Trionychinae and Cyclanorbinae, which is consistent with morphological data and molecular analysis. Our phylogenetic analyses have established a sister taxon relationship between the Asian Rafetus and the Asian Palea + Pelodiscus + Dogania + Nilssonia + Amyda, whereas a previous study grouped the Asian Rafetus with the American Apalone. The results of divergence time estimates and area ancestral reconstruction show that extant Trionychidae originated in Asia at around 108 million years ago (MA), and radiations mainly occurred during two warm periods, namely Late Cretaceous–Early Eocene and Oligocene. By combining the estimated divergence time and the reconstructed ancestral area of softshell turtles, we determined that the dispersal of softshell turtles out of Asia may have taken three routes. Furthermore, the times of dispersal seem to be in agreement with the time of the India–Asia collision and opening of the Bering Strait, which provide evidence for the accuracy of our estimation of divergence time. Overall, the mitogenomes of this group were used to explore the origin and dispersal route of Trionychidae and have provided new insights on the evolution of this group.     

Turtle origins: insights from phylogenetic retrofitting and molecular scaffolds

Adding new taxa to morphological phylogenetic analyses without substantially revising the set of included characters is a common practice, with drawbacks (undersampling of relevant characters) and potential benefits (character selection is not biased by preconceptions over the affinities of the ‘retrofitted’ taxon). Retrofitting turtles (Testudines) and other taxa to recent reptile phylogenies consistently places turtles with anapsid‐grade parareptiles (especially Eunotosaurus and/or pareiasauromorphs), under both Bayesian and parsimony analyses. This morphological evidence for turtle–parareptile affinities appears to contradict the robust genomic evidence that extant (living) turtles are nested within diapsids as sister to extant archosaurs (birds and crocodilians). However, the morphological data are almost equally consistent with a turtle–archosaur clade: enforcing this molecular scaffold onto the morphological data does not greatly increase tree length (parsimony) or reduce likelihood (Bayesian inference). Moreover, under certain analytic conditions, Eunotosaurus groups with turtles and thus also falls within the turtle–archosaur clade. This result raises the possibility that turtles could simultaneously be most closely related to a taxon traditionally considered a parareptile (Eunotosaurus) and still have archosaurs as their closest extant sister group.     

Multiple data sets, high homoplasy, and the phylogeny of softshell turtles (Testudines: Trionychidae)

We present a phylogenetic hypothesis and novel, rank-free classification for all extant species of softshell turtles (Testudines:Trionychidae). Our data set included DNA sequence data from two mitochondrial protein-coding genes and a approximately 1-kb nuclear intron for 23 of 26 recognized species, and 59 previously published morphological characters for a complimentary set of 24 species. The combined data set provided complete taxonomic coverage for this globally distributed clade of turtles, with incomplete data for a few taxa. Although our taxonomic sampling is complete, most of the modern taxa are representatives of old and very divergent lineages. Thus, due to biological realities, our sampling consists of one or a few representatives of several ancient lineages across a relatively deep phylogenetic tree. Our analyses of the combined data set converge on a set of well-supported relationships, which is in accord with many aspects of traditional softshell systematics including the monophyly of the Cyclanorbinae and Trionychinae. However, our results conflict with other aspects of current taxonomy and indicate that most of the currently recognized tribes are not monophyletic. We use this strong estimate of the phylogeny of softshell turtles for two purposes: (1) as the basis for a novel rank-free classification, and (2) to retrospectively examine strategies for analyzing highly homoplasious mtDNA data in deep phylogenetic problems where increased taxon sampling is not an option. Weeded and weighted parsimony, and model-based techniques, generally improved the phylogenetic performance of highly homoplasious mtDNA sequences, but no single strategy completely mitigated the problems of associated with these highly homoplasious data. Many deep nodes in the softshell turtle phylogeny were confidently recovered only after the addition of largely nonhomoplasious data from the nuclear intron.     

广州市花地湾市场龟鳖类调查

通过对广州市花地湾花鸟鱼虫市场贸易龟鳖类进行为期3年27次调查,共记录了9科29属41种或亚种,包括鳖科(Trionychidae)2属2种,两爪鳖科(Carettochelyidae)1属1种,龟科(Emydidae)4属5种,淡水龟科(Geoemydidae)10属18种,侧颈龟科(Chelidae)3属3种,动胸龟科(Kinosternidae)2属3种,鳄龟科(Chelydridae)2属2种,陆龟科(Testudinidae)4属6种,海龟科(Cheloniidae)1属1种。其中,国内龟鳖类共13种,外来物种共28种。受保护种类30种,所占比例达73.17%,部分龟类来源为野外捕捉。大量实体商家转入互联网进行精确营销,尤其是针对高价值的濒危种类,未来将着重对该隐蔽领域进行深入调查。     

The cranial arteries of turtles and their evolutionary significance

In this paper the cranial arteries, cranial arterial foramina, and bony canals of the Cheloniidae, Chelydridae, Pelomedusidae, and Chelidae are described in detail. From skull studies and published material, the general cranial arterial patterns of all the turtle families can be inferred. Sea turtles, the Cheloniidae and Dermochelyidae, possess both a large stapedial artery and a large artery supplying the orbit, which is possibly similar to the primitive cranial arterial pattern for turtles. From a primitive pattern in which stapedial and palatine arteries supply the orbit, the Chelydridae and Testudinidae retained a large stapedial artery and reduced the palatine artery, while the Kinosternidae and Dermatemydidae developed a large palatine artery and reduced the stapedial artery. The Trionychidae and probably the Carettochelyidae evolved a complex arterial pattern in which the stapedial artery was reduced somewhat and the pseudopalatine artery was substituted for the palatine artery. Pleurodires in general retained a large stapedial artery and reduced or eliminated the palatine artery. The Podocneminae, including the Madagascar species, developed a highly modified carotid canal, which is found in no other turtle group. The facts which have been presented should aid in fossil skull studies and in understanding the evolutionary background of recent turtles.     

Evolutionary and developmental aspects of phalangeal formula variation in pig-nose and soft-shelled turtles (Carettochelyidae and Trionychidae)

In order to examine the evolution of the phalangeal formula in a diverse clade of turtles, including hyperphalangy as a rare condition in this group, we studied 210 specimens representing all extant genera of Trionychidae and their sister taxon, Carettochelyidae. Both groups consist of highly aquatic species with elongated autopods that are either paddle-like (Trionychidae) or transformed to flippers (Carettochelyidae). Phalangeal formulae were obtained mostly by radiographs of alcohol-preserved or dry specimens, as well as by direct counts from skeletons. All trionychids and Carettochelys are pentadactylous, but their phalangeal formulae differ. Carettochelys exhibits the turtle-plesiomorphic state (manus and pes: 2-3-3-3-3), with no variation in adults. Trionychids exhibit intraspecific variation, ranging from 2-3-3-3-2 to 2-3-3-6-5 for the manus, and from 2-3-3-3-2 to 2-3-3-5-3 for the pes. The extant Carettochelys as well as the Middle Eocene Allaeochelys crassesculpta are characterized by an elongation of phalanges, whereas trionychids consistently have shorter phalanges. All trionychid genera exhibit some degree of hyperphalangy in digits IV and V, in both the manus and pes. Phalanges of the clawed digits I–III are very robust compared to phalanges of the non-clawed digits IV and V. The latter contribute significantly to the enlargement of the paddle by their additional phalanges. We hypothesize that this phalangeal pattern is coupled with prolongation of growth processes in the non-clawed digits. The differences in autopod morphology between carettochelyids and trionychids reflect different locomotor patterns related to different natural histories (elongated flippers for high-speed escape in the mainly herbivorous Carettochelys; broad paddles for rapid turns during hunting in the mainly carnivorous trionychids). The autopod of Pelodiscus sinensis is proposed as an experimental model to examine the developmental basis of adult autopod variation.     

MicroRNAs support a turtle + lizard clade

Despite much interest in amniote systematics, the origin of turtles remains elusive. Traditional morphological phylogenetic analyses place turtles outside Diapsida-amniotes whose ancestor had two fenestrae in the temporal region of the skull (among the living forms the tuatara, lizards, birds and crocodilians)-and allied with some unfenestrate-skulled (anapsid) taxa. Nonetheless, some morphological analyses place turtles within Diapsida, allied with Lepidosauria (tuatara and lizards). Most molecular studies agree that turtles are diapsids, but rather than allying them with lepidosaurs, instead place turtles near or within Archosauria (crocodilians and birds). Thus, three basic phylogenetic positions for turtles with respect to extant Diapsida are currently debated: (i) sister to Diapsida, (ii) sister to Lepidosauria, or (iii) sister to, or within, Archosauria. Interestingly, although these three alternatives are consistent with a single unrooted four-taxon tree for extant reptiles, they differ with respect to the position of the root. Here, we apply a novel molecular dataset, the presence versus absence of specific microRNAs, to the problem of the phylogenetic position of turtles and the root of the reptilian tree, and find that this dataset unambiguously supports a turtle + lepidosaur group. We find that turtles and lizards share four unique miRNA gene families that are not found in any other organisms' genome or small RNA library, and no miRNAs are found in all diapsids but not turtles, or in turtles and archosaurs but not in lizards. The concordance between our result and some morphological analyses suggests that there have been numerous morphological convergences and reversals in reptile phylogeny, including the loss of temporal fenestrae.     

The buccopharyngeal mucosa of the turtles (testudines)

Gross and histological examination of all extant families of turtles revealed that the buccopharyngeal mucosa is morphologically highly varied. The tongues of aquatic species have small lingual papillae or lack them entirely, while terrestrial species have tongues with numerous glandular papillae. The pharynx and the esophagus also have papillae in some species. These either facilitate swallowing in which case they are long, pointed, keratinized, and occur commonly in marine turtles, or they are vascular and nonkeratinized, facilitate respiratory gas exchange and are found in the Trionychidae, Dermatemyidae, and Carettochelyidae. The morphology of the buccopharyngeal mucosa of turtles reflects their diet, feeding behavior, habitat, and relationships. Convergence in the morphology of the buccopharyngeal mucosa occurs among families, especially among the Emydidae and other familes of turtles. Intergeneric parallelism is also seen within the Emydidae.     

Interordinal relationships of birds and other reptiles based on whole mitochondrial genomes

Several different groups of birds have been proposed as being the oldest or earliest diverging extant lineage within the avian phylogenetic tree, particularly ratites (Struthioniformes), waterfowl (Anseriformes), and shorebirds (Charadriiformes). Difficulty in resolving this issue stems from several factors, including the relatively rapid radiation of primary (ordinal) bird lineages and the lack of characters from an extant outgroup for birds that is closely related to them by measure of time. To help resolve this question, we have sequenced entire mitochondrial genomes for five birds (a rhea, a duck, a falcon, and two perching birds), one crocodilian, and one turtle. Maximum parsimony and maximum likelihood analyses of these new sequences together with published sequences (18 taxa total) yield the same optimal tree topology, in which a perching bird (Passeriformes) is sister to all the other bird taxa. A basal position for waterfowl among the bird study taxa is rejected by maximum likelihood analyses. However, neither the conventional view, in which ratites (including rhea) are basal to other birds, nor tree topologies with falcon or chicken basal among birds could be rejected in the same manner. In likelihood analyses of a subset of seven birds, alligator, and turtle (9 taxa total), we find that increasing the number of parameters in the model shifts the optimal topology from one with a perching bird basal among birds to the conventional view with ratites diverging basally; moreover, likelihood scores for the two trees are not significantly different. Thus, although our largest set of taxa and characters supports a tree with perching birds diverging basally among birds, the position of this earliest divergence among birds appears unstable. Our analyses indicate a sister relationship between a waterfowl/chicken clade and ratites, relative to perching birds and falcon. We find support for a sister relationship between turtles and a bird/crocodilian clade, and for rejecting both the Haemothermia hypothesis (birds and mammals as sister taxa) and the placement of turtles as basal within the phylogenetic tree for amniote animals.     

Phylogeny of the Falconidae (Aves): a comparison of the efficacy of morphological, mitochondrial, and nuclear data

We sequenced 2800+ bp of the RAG-1 exon for representatives of all the currently recognized genera in the avian family Falconidae. A phylogenetic analysis of these data was compared to prior analyses of mitochondrial (cytochrome-b) and morphological (syringeal) data. The nuclear RAG-1 sequences produced results that were in agreement with the morphological results, but differed from the mitochondrial results with regard to monophyly of the genus Micrastur. A reanalysis of the cytochrome-b (cyt-b) data suggested that this result was due to heterogeneity in base composition. Comparisons of data quality and quantity across the three data sets indicate that the nuclear DNA sequences and the morphological data have similar consistency and retention indices as well as noise distributions that are superior to those of cyt-b. However, the RAG-1 data identify more nodes with high bootstrap support indices than do either morphology or mitochondrial sequences. In the final assessment, RAG-1 sequences were superior in phylogenetic utility both to syringeal morphology (because of sheer number of characters) and to cyt-b sequences (because of reduced noise and homogeneity of base composition, but in spite of having many fewer characters).     

Comparison research and phylogenetic implications of mitochondrial control regions in four soft-shelled turtles of Trionychia (Reptilia, Testudinata)

The mitochondrial control regions (CRs) and flanking sequences of Pelodiscus sinensis, Apalone ferox, Palea steindachneri and Carettochelys insculpta were obtained using Long-PCR with gene-specific primers. The CR lengths of the four species were 1843 bp, 1356 bp, 1725 bp, and 969 bp. The base composition percentages of A+T were 60.5%, 60.7%, 65.7%, 64.7%, respectively. Combined with CR sequences of other three soft-shelled turtles published in GenBank (Pelodiscus sinensis, Korea, AY962573; Dogania subplana, AF366350; Lissemys punctata, EF050073), we compared the CR structures and identified three functional domains (TAS, CD and CSB) in which conserved sequence blocks (TAS, CSB -F, CSB-1, CSB-2 and CSB-3) were also successfully identified according to their sequence similarities to those of other turtles. The variable numbers of tandem repeats (VNTRs 1) with 50–52 bp motif were identified at 5′-end of CR among the five soft-shelled turtles P. sinensis (China), P. sinensis (Korea), A. ferox, P. steindachneri, D. subplana. The copy number of the VNTRs varied from 5 to 15. VNTRs 2 with 2–11 bp motif were identified in the 3′- end of CR among all of the six soft-shelled turtles with variable number of motifs from 4 to 29. Moreover, VNTRs 3 with 6 bp motif were identified between CSB-1 and CSB-2 of CR both in P. sinensis (China) and P. sinensis (Korea), in which the number of motifs varied from 19 to 29. The types and distribution of VNTRs of the six soft-shelled turtles were also discussed. With Alligator mississippiensis as an outgroup, combined with the CR sequences (excluding VNTRs) of other five turtles which were published in GenBank, the molecular phylogenetic trees were constructed using PAUP 4.0b10 and MrBayes ver. 3.0. The results strongly supported the monophyly of Carretochelyidae and Carettochelyidae as sister group to an assemblage of Cryptodira. Our research suggested that the earliest phylogenetic tree splits into three separated basal branches; the Pelomedusidira (Pelomedusa subrufa), the Carettochelyidae (C. insculpta), and an assemblage of Cryptodira and the C. insculpta that might be a representation of distinctive suborder.     

Phylogenetic hypotheses for the turtle family Geoemydidae

The turtle family Geoemydidae represents the largest, most diverse, and most poorly understood family of turtles. Little is known about this group, including intrafamilial systematics. The only complete phylogenetic hypothesis for this family positions geoemydids as paraphyletic with respect to tortoises, but this arrangement has not been accepted by many workers. We compiled a 79-taxon mitochondrial and nuclear DNA data set to reconstruct phylogenetic relationships for 65 species and subspecies representing all 23 genera of the Geoemydidae. Maximum parsimony (MP) and maximum-likelihood (ML) analyses and Bayesian analysis produced similar, well-resolved trees. Our analyses identified three main clades comprising the tortoises (Testudinidae), the old-world Geoemydidae, and the South American geoemydid genus Rhinoclemmys. Within Geoemydidae, many nodes were strongly supported, particularly based on Bayesian posterior probabilities of the combined three-gene dataset. We found that adding data for a subset of taxa improved resolution of some deeper nodes in the tree. Several strongly supported groupings within the Geoemydidae demonstrate non-monophyly of some genera and possible interspecific hybrids, and we recommend several taxonomic revisions based on available evidence.     

Complete mitochondrial DNA sequences of the green turtle and blue-tailed mole skink: statistical evidence for archosaurian affinity of turtles.

Turtles have highly specialized morphological characteristics, and their phylogenetic position has been under intensive debate. Previous molecular studies have not established a consistent and statistically well supported conclusion on this issue. In order to address this, complete mitochondrial DNA sequences were determined for the green turtle and the blue-tailed mole skink. These genomes possess an organization of genes which is typical of most other vertebrates, such as placental mammals, a frog, and bony fishes, but distinct from organizations of alligators and snakes. Molecular evolutionary rates of mitochondrial protein sequences appear to vary considerably among major reptilian lineages, with relatively rapid rates for snake and crocodilian lineages but slow rates for turtle and lizard lineages. In spite of this rate heterogeneity, phylogenetic analyses using amino acid sequences of 12 mitochondrial proteins reliably established the Archosauria (birds and crocodilians) and Lepidosauria (lizards and snakes) clades postulated from previous morphological studies. The phylogenetic analyses further suggested that turtles are a sister group of the archosaurs, and this untraditional relationship was provided with strong statistical evidence by both the bootstrap and the Kishino-Hasegawa tests. This is the first statistically significant molecular phylogeny on the placement of turtles relative to the archosaurs and lepidosaurs. It is therefore likely that turtles originated from a Permian-Triassic archosauromorph ancestor with two pairs of temporal fenestrae behind the skull orbit that were subsequently lost. The traditional classification of turtles in the Anapsida may thus need to be reconsidered.     

A neglected lineage of North American turtles fills a major gap in the fossil record

Abstract: The fossil record of the two primary subclades of softshell turtles (Trionychidae) is exceedingly asymmetric, as a result of a ghost range of total clade Cyclanorbinae that is estimated at 80 Ma. Herein, we present the first phylogenetic analysis of Trionychidae that includes a representative of the poorly studied taxon Plastomenidae, which is known from the Campanian to Eocene of North America. The analysis reveals that plastomenids are stem cyclanorbines, thus significantly reducing the apparent ghost range of total group Cyclanorbinae to approximately 30 Ma. Plastomenids are either an early branching clade of stem Cyclanorbinae, or they represent a paraphyletic grade that gave rise to modern cyclanorbines. Although abundant, the fossil record is still too poorly understood to distinguish between these two primary hypotheses. The previously persistent extremely long ghost range of total clade Cyclanorbinae appears to have been the result of a research bias.     

Proopiomelanocortin (POMC) and testing the phylogenetic position of turtles (Testudines)

Morphological and molecular studies have inferred multiple hypotheses for the phylogenetic relationships of Testudines. The hypothesis that Testudines are the only extant anapsid amniotes and the sister taxon of diapsid amniotes is corroborated by morphological studies, while the hypothesis that Testudines are diapsid amniotes is corroborated by more recent molecular and morphological studies. In this study, the placement of Testudines is tested using the full length cDNA sequence of the polypeptide hormone precursor proopiomelanocortin (POMC). Because only extant taxa have been used, the hypotheses being tested are limited to the following (1) Testudines as the sister taxon of Archosauria, (2) Testudines included in Archosauria and the sister taxon of Crocodilia, (3) Testudines as the sister taxon of Lepidosauria, (4) Testudines as the sister taxon of Sauria, and (5) Testudines as the sister taxon of a monophyletic Mammalia–Sauria clade. Neither Maximum likelihood, Bayesian, or maximum parsimony analyses are able to falsify the hypothesis of (Archosauria (Lepidosauria, Testudines)) and as such is the preferred inference from the POMC data.     

A Phylogenomic Approach to Vertebrate Phylogeny Supports a Turtle-Archosaur Affinity and a Possible Paraphyletic Lissamphibia

In resolving the vertebrate tree of life, two fundamental questions remain: 1) what is the phylogenetic position of turtles within amniotes, and 2) what are the relationships between the three major lissamphibian (extant amphibian) groups? These relationships have historically been difficult to resolve, with five different hypotheses proposed for turtle placement, and four proposed branching patterns within Lissamphibia. We compiled a large cDNA/EST dataset for vertebrates (75 genes for 129 taxa) to address these outstanding questions. Gene-specific phylogenetic analyses revealed a great deal of variation in preferred topology, resulting in topologically ambiguous conclusions from the combined dataset. Due to consistent preferences for the same divergent topologies across genes, we suspected systematic phylogenetic error as a cause of some variation. Accordingly, we developed and tested a novel statistical method that identifies sites that have a high probability of containing biased signal for a specific phylogenetic relationship. After removing putatively biased sites, support emerged for a sister relationship between turtles and either crocodilians or archosaurs, as well as for a caecilian-salamander sister relationship within Lissamphibia, with Lissamphibia potentially paraphyletic.     

中华鳖线粒体基因组序列分析

参照近源物种线粒体基因组序列,设计17对特异引物,采用PCR产物直接测序法测得中华鳖线粒体基因组全序列.初步分析其基因组特点和各基因的定位,用pDRAW32软件预测12种限制性酶对其的酶切图谱.结果表明,中华鳖线粒体基因组全长17364bp,核苷酸组成为35.23%A、27.26%T、25.73%C、11.78%G,包括13个蛋白质编码基因、2个rRNA基因、22个tRNA基因和1个非编码控制区.基于线粒体基因组编码的13个蛋白质的氨基酸序列,用NJ法和MP法构建系统进化树,分析6种龟鳖类动物之间的亲缘关系,与传统的系统分类基本一致,初步确定淡水龟科与海龟科的亲缘关系比与龟科的亲缘关系要近.     

The genus Coleodactylus (Sphaerodactylinae, Gekkota) revisited: a molecular phylogenetic perspective

Nucleotide sequence data from a mitochondrial gene (16S) and two nuclear genes (c-mos, RAG-1) were used to evaluate the monophyly of the genus Coleodactylus, to provide the first phylogenetic hypothesis of relationships among its species in a cladistic framework, and to estimate the relative timing of species divergences. Maximum Parsimony, Maximum Likelihood and Bayesian analyses of the combined data sets retrieved Coleodactylus as a monophyletic genus, although weakly supported. Species were recovered as two genetically and morphological distinct clades, with C. amazonicus populations forming the sister taxon to the meridionalis group (C. brachystoma, C. meridionalis, C. natalensis, and C. septentrionalis). Within this group, C. septentrionalis was placed as the sister taxon to a clade comprising the rest of the species, C. meridionalis was recovered as the sister species to C. brachystoma, and C. natalensis was found nested within C. meridionalis. Divergence time estimates based on penalized likelihood and Bayesian dating methods do not support the previous hypothesis based on the Quaternary rain forest fragmentation model proposed to explain the diversification of the genus. The basal cladogenic event between major lineages of Coleodactylus was estimated to have occurred in the late Cretaceous (72.6+/-1.77 Mya), approximately at the same point in time than the other genera of Sphaerodactylinae diverged from each other. Within the meridionalis group, the split between C. septentrionalis and C. brachystoma+C. meridionalis was placed in the Eocene (46.4+/-4.22 Mya), and the divergence between C. brachystoma and C. meridionalis was estimated to have occurred in the Oligocene (29.3+/-4.33 Mya). Most intraspecific cladogenesis occurred through Miocene to Pliocene, and only for two conspecific samples and for C. natalensis could a Quaternary differentiation be assumed (1.9+/-1.3 Mya).     

A survey of the epibiota of Eretmochelys imbricata (Testudines: Cheloniidae) of Mona Island, Puerto Rico

Epibiotic organisms inhabiting non-nesting hawksbill sea turtles, Eretmochelys imbricata (Linnaeus, 1766), are described from Mona and Monito Islands, Puerto Rico. Epibiont samples from 105 turtles of shallow (< 40 m) water foraging habitats were collected and identified to the lowest possible taxon. This epibiotic assemblage consisting of at least 4 algal functional groups and 12 animal phyla represents the greatest phylogenetic diversity for marine turtle epibiota. Six groups are considered new reports for marine turtles. Most epibiont colonization was found on posterior marginal scutes and under overlapping scutes. Ecological attributes of epibiota and their symbiosis with E. imbricata provide a tool to understand basi and epibiont populations.