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Bonita mexicana new genus, new species, Point Sofia, Tortugas Bay, Baja California Sur, México (CNCR 25300). Adult female, abdomen and telson: A, C, outer surface; B, inner surface. Scales: A = 2.76 mm; B = 2.64 mm; C = 3.47 mm.
Source publication
Two monotypic genera, Enigmatheres new genus, and Bonita new genus, are described to receive Fabia canfieldi Rathbun, 1918, and Bonita mexicana new species respectively. Enigmatheres can be distinguished from other Pinnotheridae by the third maxilliped having a gently curved outer margin, with a carpus that is larger than the conical propodus and a...
Contexts in source publication
Context 1
... greatly enlarged, with 6 somites and telson well separated (Fig. 4 A), strongly concave, laterally covering ischia, margin with tomentum, involuted, (Fig. 4B-C), distally covering the buccal cavity. Male. Unknown. Etymology. The name mexicana refers to the country where the type material was collected. Remarks. The soft carapace, the protuberance on the basal article of the antennae, the transversely ...
Context 2
... greatly enlarged, with 6 somites and telson well separated (Fig. 4 A), strongly concave, laterally covering ischia, margin with tomentum, involuted, (Fig. 4B-C), distally covering the buccal cavity. Male. Unknown. Etymology. The name mexicana refers to the country where the type material was collected. Remarks. The soft carapace, the protuberance on the basal article of the antennae, the transversely placed MXP3 on the buccal cavity, the slender and feeble walking legs and the well-separated ...
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Citations
... Some pea crab species have been well-studied in terms of reproduction (Hines 1992;Bolaños et al. 2004;Becker and Türkay 2010;Campos 2013;Peiró et al. 2013;de Gier and Becker 2020) and host use (Schmitt et al. 1973;Palacios Theil et al. 2016;de Gier and Becker 2020). Pea crab taxonomy has undergone many recent revisions-based both on shared morphological traits and phylogenetic studies (Harrison 2004;Ocampo et al. 2013;Palacios Theil et al. 2016)-such as the recent examination of the subfamily Pinnotherinae (Campos 2009) or the establishment of the subfamily Pinnixulalinae (Palacios Theil et al. 2016). Importantly, molecular phylogenetics has revealed that some pinnotherid genera, originally defined by shared morphological traits, are paraphyletic, a trend thought to result from convergent morphological evolution to different hosts (Palacios Theil et al. 2016). ...
... Although the intent of our study was not to test taxonomic hypotheses for the group, the phylogeny we constructed-together with host use patterns-correspond to some (but not all) of the recent taxonomic and phylogenetic revisions proposed for this group (Campos 2009;Palacios Theil et al. 2016;Palacios Theil and Felder 2020). For example, the subfamily Pinnotherinae are characterized morphologically by a soft carapace and slender, feeble walking legs (Campos 2009). ...
... Although the intent of our study was not to test taxonomic hypotheses for the group, the phylogeny we constructed-together with host use patterns-correspond to some (but not all) of the recent taxonomic and phylogenetic revisions proposed for this group (Campos 2009;Palacios Theil et al. 2016;Palacios Theil and Felder 2020). For example, the subfamily Pinnotherinae are characterized morphologically by a soft carapace and slender, feeble walking legs (Campos 2009). While our phylogeny does not resolve the Pinnotherinae as a monophyletic group (Fig. 2; Pinnaxodes to Afropinnotheres), it does resolve the two major clades in the Pinnotherinae recovered by recent phylogenies (Palacios Theil et al. 2016), and demonstrates that they share a similar body shape (square or circular) and similar set of hosts (bivalves, tunicates, limpets, and echinoderms). ...
Many marine crustaceans form symbiotic relationships, yet there has been little work quantifying morphological adaptations in species specialized on different hosts. Here we examine morphological adaptations of symbiotic pea crabs (Pinnotheridae) to different host phyla. Multiple authors have noted that crabs living with burrowing hosts (burrowing shrimps and annelids) have different carapace shapes than species living with non-burrowing hosts (bivalves, gastropods, echinoderms, and ascidians), but this hypothesis has never been tested. Using digital analyses of taxonomic images, we calculated carapace aspect ratio of 149 pinnotherid species, and used phylogenetic ANOVA to test whether aspect ratio differed among species living with different hosts. Pea crab species living with burrowing hosts had significantly larger carapace aspect ratios (wide carapaces) than species living with non-burrowing hosts (round or square carapaces, or otherwise non-wider than long). Convergent evolution of morphological features in species with specialized host use may be a common—but understudied—pattern in many marine groups.
... Although the functionality is unknown, species of some pea crab genera possess an elongated rostrum, like Austrotheres [12], Limotheres [55], Serenotheres (e.g., [27]) and, to a lesser degree, in Abyssotheres [60] and Nepinnotheres (e.g., [7]). Members of the (paraphyletic) genus Fabia and the related genus Bonita possess an extension of the rostrum towards the midline of the carapace: two longitudinal sulci split the anterior side of the carapace in three portions [16,61]. ...
... longitudinal sulci split the anterior side of the carapace in three portions [16,61]. ...
... A common associated feature with such elongated chelae is a setal ornamentation of the inner surface of the palm and pollex. This brush-like row of setae can be found in female specimens of many genera associated with bivalves: Abyssotheres, Afropinnotheres, Amusiotheres, Arcotheres, Austrotheres, Bonita, Fabia ( Figure 6C), Gemmotheres, Discorsotheres, Durckheimia, Latatheres, Nannotheres, Nepinnotheres (but not N. pinnotheres), Pinnotheres ( Figure 6D), Sindheres, Tacitotheres, Viridotheres, Visayeres, Xanthasia, Waldotheres, and Zaops [7,12,15,16,19,27,30,31,54,60,61,[102][103][104][105]. This adaptive feature can also be found in two genera associated with gastropods, Ernestotheres and Calyptraeotheres [7,89], and in the sea urchin-associated Dissodactylus latus Griffith, 1987 [93]. ...
Almost all pea crab species in the subfamily Pinnotherinae (Decapoda: Brachyura: Pinnotheridae) are considered obligatory endo-or ectosymbionts, living in a mutualistic or parasitic relationship with a wide variety of invertebrate hosts, including bivalves, gastropods, echinoids, holothurians, and ascidians. While the subfamily is regarded as one of the most morphologically adapted groups of symbiotic crabs, the functionality of these adaptations in relation to their lifestyles has not been reviewed before. Available information on the ecomorphological adaptations of various pinnotherine crab species and their functionality was compiled in order to clarify their ecological diversity. These include the size, shape, and ornamentations of the carapace, the frontal appendages and mouthparts, the cheliped morphology, the ambulatory legs, and the reproductive anatomy and larval characters. The phylogenetic relevance of the adaptations is also reviewed and suggestions for future studies are made. Based on an updated list of all known pinnotherine symbiont-host associations and the available phylogenetic reconstructions, it is concluded that, due to convergent evolution, unrelated species with a similar host interaction might display the same morphological adaptations.
... The symbiotic way of life, sexual dimorphism, and small size of these crabs has complicated their taxonomic classification (Becker and Türkay, 2010). The systematics of pinnotherids has been the subject of numerous revisions at different taxonomic levels (Rathbun, 1918;Balss, 1957;Manning, 1993;Ahyong and Ng, 2009;Campos, 2009;Becker and Türkay, 2010;Palacios Theil et al., 2016). These revisions have been based mainly on adult morphology. ...
Pinnotheridae comprises a diverse group of small crabs that live as symbionts with invertebrates. All Pinnotheridae are marine species with a worldwide distribution, living in close relationship with their hosts. Independently of the type of host or symbiotic relationship (parasitic, commensalistic), all species with known larval development present free-living larval stages (except Tunicotheres moseri, where larval stages develop in the abdominal enclosure of the parental female). The larval development comprises a zoea phase, with a variable number of stages (2–5), and a megalopa phase with a single stage. Approximately 290 pinnotherid species have been described, and information on larval developmental stages is available for 54 of them. This leaflet presents the distinctive features of the larval stages for 12 out of 13 species distributed in ICES area: Pinnixulala retinens, Pinnixa cylindrica, P. lunzi, Rathbunixa sayana, Tubicolixa chaetopterana, Afropinnotheres monodi, Dissodactylus mellitae, Nepinnotheres pinnotheres, Pinnotheres bicristatus, P. pectunculi, P. pisum, Tumidotheres maculatus, and Zaops ostreus. Complete larval development is only known for 6 of these species. Illustrated keys to identify the known zoea and megalopa stages are included.
... Because other pinnotherid crabs may infest two or more hosts (Campos, 2016;Hernández et al., 2017), we believe the presence of G. chamae in other species of Chama would be possible, so we recommend examining additional Chama species throughout its range. Other pinnotherid crab species symbiotic with Jewel Box clams include Bonita mexicana Campos, 2009 (Pacific, Baja California, México in Pseudochama exogyra (Conrad, 1837)) and Durckheimia caeca Bürger, 1895(Palau and Japan, in Ch. reflexa Reeve, 1846[= Ch. pacifica Broderip, 1835) (Ahyong and Ng, 2005;Campos, 2009). ...
... Because other pinnotherid crabs may infest two or more hosts (Campos, 2016;Hernández et al., 2017), we believe the presence of G. chamae in other species of Chama would be possible, so we recommend examining additional Chama species throughout its range. Other pinnotherid crab species symbiotic with Jewel Box clams include Bonita mexicana Campos, 2009 (Pacific, Baja California, México in Pseudochama exogyra (Conrad, 1837)) and Durckheimia caeca Bürger, 1895(Palau and Japan, in Ch. reflexa Reeve, 1846[= Ch. pacifica Broderip, 1835) (Ahyong and Ng, 2005;Campos, 2009). ...
... Remarks. Campos (2009) included Gemmotheres within the Pinnotherinae sensu stricto, the principal synapomorphies of the adult female phase being a soft, thin carapace, and a subconical protuberance on the basal article of the antenna (Fig. 1B). This protuberance is, in fact, an extension of article 1, which has the nephridiopore of the antennal gland (= green gland) (Davie et al., 2015;Ng et al., 2019). ...
An adult female of the Jewel-Box clam crab Gemmotheres chamae (Roberts, 1975) was collected from a shrimp trawl during a biological exploration off Campeche coast, Mexico. This finding represents its first record in the Gulf of Mexico and the second locality for this species along the Atlantic coast of America, the first being from off North Carolina, U.S.A. As an adult, G. chamae has a soft, thin carapace, and a subconical protuberance on article 1 of the antennae (with the nephridiopore of the antennal gland), so it is considered to belong to the Pinnotherinae sensu stricto. G. chamae and Nannotheres moorei Manning and Felder, 1996 (Atlantic) are the only members of the American Pinnotherinae sensu stricto that have the maxilliped 3 with a 2-segmented palp. The asymmetry of pereiopod 3 is confirmed, as well as that of the pereiopod 4, but the right legs are the longest. All these features are diagnostic for G. chamae.
... However, among a wider suite of genera that we have assessed for the form of the antennal articles, including Alain Manning, 1998 (see also Ahyong & Ng, 2008), Buergeres Ng & Manning, 2003, Pinnotheres Bosc, 1802, Nepinnotheres Manning, 1993a, Arcotheres Manning, 1993a, Holotheres Ng & Manning, 2003, Viridotheres Manning, 1996, Serenotheres Ahyong & Ng, 2005, and Visayeres Ahyong & Ng, 2007b, no pattern in relation to somatic morphology or host is yet evident. We note that the "basal antennal article" referred to by Campos ( , 2009) represents antennal article 1 and is not equivalent to the "basal antennal article" frequently used in brachyuran nomenclature, especially in the taxonomy of Majoidea (Davie et al., 2015: 64). The "basal antennal article" in more common usage among brachyurans, rather than referring to article 1, refers to the quadrate or subquadrate structure just below the orbit, which is composed of the fused articles 2 and 3. Antennal article 1 is the osmoregulatory article, which contains the green gland and its opening, the nephridiopore. ...
... In many crabs, article 1 is often small or even indistinguishably fused to the surface of the epistome, but the presence of the nephridiopore indicates its position. In all the pinnotherids examined (see also Campos, , 2009Ahyong & Ng, 2005, 2007a, b, 2008Ng & Meyer, 2016), the fused articles 2 and 3 are usually demarcated by shallow grooves and article 1 is also immovably fused but positioned very low on the epistome such that its lower margin intrudes into, and forms part of, the margin of the buccal cavity. This lower margin of article 1 varies from indistinct to bluntly angular, appearing as a short lobe or protuberance, being most pronounced mesially. ...
... This mesioventral position of the presumed nephridiopore, readily observed in larger specimens of Nepinnotheres and Arcotheres, this may well be the normal position in most, if not all, pinnotherines (and pinnotherids). Using primarily molecular data, Palacios-Theil et al. (2016) recognised a Pinnotherinae (based mostly on American representatives, including Mesotheres barbatus), somewhat different from the system proposed by Campos (2009) which emphasised morphological characters like the presence or absence of a protuberance on the inner ventral edge of antennal article 1 as well as known larval features. The location of antennal article 1, on the margin of the buccal cavity, may prove to be a synapomorphy of Pinnotheridae, but until more comprehensive phylogenetic and comparative morphological studies can be completed, the taxonomic and phylogenetic value of this character remains unclear. ...
Orthotheres Sakai, 1969, was recently restricted to three species from the western Pacific, noting that other species assigned to this genus from the Americas, Philippines and Palau belonged to other genera. Studies of the non-Orthotheres material confirm that the species from the Americas should be placed in a new genus, Mesotheres, characterised by the structure of the carapace, eyes, third maxilliped and ambulatory legs. Mesotheres includes Pinnotheres barbatus Desbonne, in Desbonne & Schramm, 1867, Pinnotheres serrei Rathbun, 1909, Pinnotheres strombi Rathbun, 1905, and Fabia unguifalcula Glassell, 1936. Three species from the western Pacific, Pinnotheres glaber Bürger, 1895, Pinnotheres laevis Bürger, 1895, and Pinnotheres longipes Bürger, 1895, are referred to a new genus, Tacitotheres, and can be distinguished by carapace form, as well as structure of the third maxilliped and ambulatory legs.
... Unpublished observations on juveniles and adults of several species of pinnotherid crabs by the author, for example, Juxtafabia muliniarum (Rathbun, 1918), Dissodactylus lockingtoni Glassell, 1935, D. xantusi Glassell, 1936, Calyptraeotheres granti (Glassell, 1933, Austinotheres angelicus (Lockington, 1877) and Tumidotheres margarita (Smith, 1870) have revealed that the third maxilliped exhibit little morphological variation through the post-larval stages of development (from juvenile to adult). This appendage, including the partial or total fusion of the ischium and merus ( Fig. 1A-G; Fig 2A, C-G), has been considered a valuable feature for distinguishing genera in the family Pinnotheridae (Bürger 1895;Rathbun 1918;Manning 1993;Ahyong & Ng 2007;Campos 2009). According to Melzer & Schwabe (2008) the ischium and merus of the third maxilliped are completely fused, the carpus is larger than the propodus and the small dactylus is subterminally inserted on the propodus (Fig 1C), and as such the juveniles studied are possibly members of the genus Orthotheres Sakai, 1969. ...
... 4); F, fromRathbun (1918: fig. 57); G, fromCampos (2009: fig 3e). Scale bars, A, C= 0.5 mm; D-E= 1.0 mm; F= 0.57 mm; G=0.23 mm. ...
The morphology of the juveniles recorded by Melzer & Schwabe (2008) supports the present hypothesis that they are not a species of Orthotheres but instead belonging to Calyptraeotheres, possibly C. politus
... Study of the taxonomy of this group is particularly difficult due to several factors: their cryptic symbiotic way of life, their small size, similar morphological adaptations for living as a symbiont, and sexual dimorphism (including different female morphologies related to maturity stage). Over the years, their taxonomy has undergone numerous revisions and changes, even recently (Ahyong and Ng 2009;Campos 2009;Palacios-Theil et al. 2016). ...
Currently, there are five species of pea crabs in
European waters. In the context of the AFROBIV project, in
which the effect of the African pea crab Afropinnotheres
monodi on bivalve species of commercial interest is being
studied, a sampling campaign has been carried out on bivalve
species of the coasts of southwestern Europe. The results of
this campaign are used in the present study to evaluate the
current distribution of A. monodi and its degree of infestation
in its various hosts, together with the distribution of native
European pinnotherid species. Previously collected information
has also been reviewed and included with the present
results to establish a baseline so as to enable future changes
in distribution and host use of European pinnotherid species to
be compared and quantified. To enable the more accurate
identification of species and detection of new species in the
area, three genetic markers (16S, Cox1, and H3) for the current
five European pinnotherid species have also been compiled
from Genbank or obtained in the present work. The results showed the high abundance of A. monodi in a significant
portion of the area studied. In contrast, few specimens of
the native Pinnotheres pectunculi and P. pisum were collected.
The northern limit of the current distribution of A. monodi
extends to the Sado estuary (Portugal), and the species has
been recorded for the first time in the Mediterranean
(Alboran Sea). New records of P. pectunculi in the Alboran
Sea confirm the presence of this species in the Mediterranean,
and considerably expand the known distribution of the
species, until now restricted to the coasts of Brittany.
... The strong and firm body of P. gigas does not match the soft carapace and weak pereiopods of pinnotherids that evolved as symbionts of bivalves, e.g. species in Pinnotherinae sensu stricto (Campos 2009). It therefore possible that geoducks clams, Panopea spp., have been more recently exploited as hosts of P. gigas. ...
New bivalve host records for four pinnotherid crabs of the Mexican Pacific are reported: Nodipecten subnodosus (Sowerby, 1835). The southernmost distribution of F. subquadrata is extended to about 600 km along the west coast of the Baja California Peninsula; the distribution of P. gigas is extended outside the Gulf of California more than 1000 km north to San Quintín, on the west coast of Baja California; and the range of T. mar-garita is restricted on the west coast of Baja California to Scammon´s Lagoon, Baja California Sur and Playa Kino Viejo, Sonora in the central region of the Gulf of California, Mexico to Panama. Based on the new material, new information on taxonomy, ecology, and life history is provided for each of these species. Pinnotheres nudus Holmes, 1895 is restored as a valid species and is removed from its synonymy with O. transversus. An updated checklist with remarks on zoogeogra-phy for the 60 pinnotherid species, included in 23 genera, of the northeastern Pacific region (Alaska to the Mexican tropical Pacific) is given.
... In the latter case, the subfamilies Anomalifrontinae Rathbun, 1931, Asthenognathinae Stimpson, 1856 and Xenophthalminae Alcock, 1900 were removed from Pinnotheridae and the species belonging to them were placed in other families on the basis of morphology or ecological characters such as swimming behaviour or the lack of typical pinnotherid symbiotic relationships (Table 1). At generic and specific levels, numerous revisions have been undertaken (for example , Rathbun 1918;Manning 1993;Ahyong and Ng 2007;Campos 2009). Yet taxonomic relationships at almost all levels remain uncertain and additional revisions are necessary (Ahyong and Ng 2007;Ahyong and Ng 2009;Campos 2009). ...
... At generic and specific levels, numerous revisions have been undertaken (for example , Rathbun 1918;Manning 1993;Ahyong and Ng 2007;Campos 2009). Yet taxonomic relationships at almost all levels remain uncertain and additional revisions are necessary (Ahyong and Ng 2007;Ahyong and Ng 2009;Campos 2009). ...
... All of the pinnotherines from Europe and Asia, in addition to some of the American species, grouped together in one of the subclades ( Fig. 1; Pinnotherinae I), whereas the second subclade was comprised exclusively of American species ( Fig. 1; Pinnotherinae II). The separation of these two clades agrees with the subdivision of Pinnotherinae proposed by Campos (2009), who divided the genera within the subfamily into two groups. One group, which he considered Pinnotherinae s.s., is characterised by a thin carapace, a protuberance on the basal antennal article, and a common generalised shape of the abdomen in the zoeal stage, all of which are absent in genera of the other group. ...
The crabs of the family Pinnotheridae are well known as commensals or parasites, mainly of molluscs and tubeworms. The phylogeny of the group, however, is poorly understood, with preliminary morphological and molecular studies questioning its monophyly. Here we used molecular genetic markers (16S, 12S mitochondrial; histone 3 nuclear) to infer a phylogeny for the family Pinnotheridae De Haan, 1833 to reevaluate the phylogeny and systematics at the level of its subfamilies and genera. Our molecular phylogeny indicated that Parapinnixa cortesi Thoma, Heard, & Vargas, 2005, Parapinnixa hendersoni Rathbun, 1918, Pinnotherelia laevigata H. Milne Edwards & Lucas, 1844, Sakaina yokoyai (Glassell, 1933), Tetrias fischerii (A. Milne-Edwards, 1867) and Tetrias scabripes Rathbun, 1898 should be removed from the family Pinnotheridae, while composition of the present subfamilies, Pinnotherinae De Haan, 1833 and Pinnothereliinae Alcock, 1900, must be revised. At generic level, Clypeasterophilus Campos, 1990, Dissodactylus Smith, 1870, Fabia Dana, 1851, Nepinnotheres Manning, 1993 and Pinnixa White, 1846 were not monophyletic in our analyses. With the exclusion of Pinnotherelia from Pinnotheridae, remaining species of Pinnothereliinae are assigned to Pinnixinae Števčić, 2005, a new subfamily based upon revision and elevation of rank for the tribe Pinnixini Števčić, 2005. In addition, we restructure membership of the subfamily Pinnotherinae and propose Pinnixulalinae, subfam. nov. to accommodate species that were excluded by molecular analyses from the other two subfamilies. These have a firm, wider-than-long carapace with clearly defined regions, strong legs that are usually tuberculate and very setose, and a third maxilliped with an elongate ischiomerus in which the ischium and merus may or may not be indistinguishably fused. Our analyses included 169 pinnotherid exemplars, representing almost half of the genera and about a quarter of the species presently recognised for the family. The relationships within and among some taxa are resolved to greater or lesser extent and the phylogenetic biodiversity of pinnotherid crabs is revealed. However, future publications will most likely result in a further increase in the number of taxa.
... Brachyuran crabs of the family Pinnotheridae in the Mexican Pacific are relatively well known, in great part due to the extensive study of this group of crabs by Ernesto Campos (see Campos 2006, 2009). The family is represented in the Mexican Pacific by 47 species (Hendrickx 1995; unpublished). ...
Calyptraeotheres camposi sp. nov. is described from the Gulf of California, Mexico. The new species is close to C. granti (Glassell, 1933) and C. pepeluisi Campos & Hernández-Ávila, 2010 from the Mexican Pacific and to C. hernandezi Hernández-Ávila & Campos 2006 from the Western Atlantic. These four species feature a third maxilliped with a 2-segmented endopod palp and the exopod with unsegmented flagellum. Calyptraeotheres camposi sp. nov. differs from C. granti and C. hernandezi by having the eyes visible in dorsal view, the carapace with arcuate anterolateral margins, the dorsal, longitudinal depressions connected with the transversal depression, and the propodus of pereiopod 2 equal or slightly longer than the carpus. From C. pepeluisi it is distinguished by the absence of a transversal depression on the carapace and the longitudinal depressions not connecting, the carpus and propodus of the third maxilliped being sub-trapezoidal and sub-conical, respectively, in lieu of subrectangular, and the inner surface of the fixed finger nude instead of bearing short setae near the cutting edge and ventral margin.
