caribboecetes progreso, a new species of sand-dwelling amphipod (amphipoda: corophiidea:...

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Zootaxa 3652 (3): 370–380 www.mapress.com/zootaxa/ Article

http://dx.doi.org/10.11646/zootaxa.3652.3.5http://zoobank.org/urn:lsid:zoobank.org:pub:3A45303E-EAF4-4299-962E-2CD745A48AF3

Caribboecetes progreso, a new species of sand-dwelling amphipod (Amphipoda: Corophiidea: Ischyroceridae) from the Gulf of Mexico, with a key for the genus

CARLOS E. PAZ-RÍOS1 & PEDRO-LUIS ARDISSON2

Departamento de Recursos del Mar, Cinvestav, Carretera antigua a Progreso, km 6, Apdo. Postal 73, Cordemex 97310 Merida, Yucatan, Mexico. E-mail: [email protected], [email protected]

Abstract

A new species of the genus Caribboecetes Just, 1983 is described and illustrated from specimens collected on sandy bot-toms of the northern Yucatan Peninsula, southeastern Gulf of Mexico. Caribboecetes progreso sp. nov. differs from the closely related species Caribboecetes barbadensis Just, 1983 and Caribboecetes jenikarpae Just, 1984 by the inflated tri-angular anterolateral flange on basis of gnathopod 2, and from Caribboecetes justi Ortiz & Lemaitre, 1997 by the setose anterolateral surface of coxal plate 7 and basis of pereopod 7. Ecological notes for the new species, a morphological com-parison, map of distribution and key for all members in the genus are also provided.

Key words: Yucatan, western Atlantic, Crustacea, Peracarida, biodiversity, taxonomy

Introduction

Members of the genus Caribboecetes Just, 1983 are currently distributed on the American coasts from the tropical western Atlantic and tropical eastern Pacific; they are commonly found on sandy or muddy bottoms in empty mollusk shells or tubes made of sand grains where they achieve high abundances (Just 1983, 1984, 1988).

The genus Caribboecetes included eight species until 1997. After that year, the genus was re-diagnosed and divided by Just (1998) to encompass five species: Caribboecetes barbadensis Just, 1983 (type species), Caribboecetes intermedius Just, 1984, Caribboecetes jenikarpae Just, 1984, Caribboecetes justi Ortiz & Lemaitre, 1997, and Caribboecetes pterycornis Just, 1984. The remaining species (Caribboecetes crassicornis Just, 1984, Caribboecetes magellani Just, 1984, and Caribboecetes squamiferus Just, 1984) were transferred to a new erected genus by Just (1998), Ambicholestes. Besides the five nominal species, the genus Caribboecetes also includes two generic species: Caribboecetes sp. sensu Just, 1984 and Caribboecetes sp. A sensu LeCroy, 2007.

The present paper describes Caribboecetes progreso sp. nov. from the Yucatan coast, southeastern Gulf of Mexico, another species inhabiting sandy bottoms in the tropical western Atlantic, increasing thus the diversity of the genus to six species. The new species is highly abundant in the Yucatan coast, nevertheless its occurrence as a new species had been overlooked. For instance, when it was found for the first time, it was misidentified asCorophium multisetosum Stock, 1952 by Ardisson and Castillo-Fernández (2004), an unrecorded species in the Gulf of Mexico so far. Later studies in the area continued reporting the new species as C. multisetosum (Rodríguez-Pliego et al. 2011; Wijnhoven et al. 2011). The present study contributes to clarify this misunderstanding.

Material and methods

The material examined herein was derived from three benthic studies in the northern Yucatan Peninsula, southeastern Gulf of Mexico. The first one was a spatio-temporal sampling along the central sector of the Yucatan State, Mexico (Ardisson & Castillo-Fernández 2004). The environmental characteristics, sampling design, sampling device and samples processing of this study are described in detail by Rodríguez-Pliego et al. (2011). To

370 Accepted by J. Lowry: 18 Apr. 2013; published: 17 May 2013


provide ecological information, along 42 km of coast and perpendicularly to the shoreline, abundance (Ind. m-2) was recorded on 12 transects 4-km apart. Along each transect, 11 sampling sites 20-m apart were fixed (from the intertidal to shallow subtidal). Four sampling campaigns were carried out: April-May 2002 (before the hurricane Isidore struck on September 2002), April-May 2003, June 2003 and November 2003. The second study consisted in a spatial sampling on the continental shelf of northern Yucatan Peninsula (Paz-Ríos & Ardisson 2013). The sampling was carried out by the Mexican Navy (Secretaría de Marina, Armada de México) during an oceanographic cruise (Yucatan-2004A, April 2004) on board the R/V Onjuko. The samples were collected with a Petite Ponar grab and fixed in 10% formalin buffered with seawater. Then they were washed in freshwater, sorted and preserved in 70% ethanol. The third study was a temporal sampling (June 2008-May 2009) in the shallow subtidal zone (~3 m depth) off a beach on the central sector of the Yucatan State, Mexico (Paz-Ríos 2009). Sampling device and samples processing in this project are the same ones described by Rodríguez-Pliego et al.(2011). In order to increase the ecological information, the distribution of abundance during an annual cycle for the new species was plotted. Data are the monthly recorded abundance (Ind. 0.25 m-2) in 12 months with five replicates for each one.

Appendages and mouthparts were mounted on glass slides with glycerol. Specimens were dissected under stereoscope microscope and illustrations were made under compound microscope with camera lucida. Gnathopods and pereopods dissected were illustrated with omitted setae in order to show clearly the whole appendages. Type material is deposited in the Colección de Invertebrados Bentónicos de Yucatán (CYMX), Cinvestav and in the Colección de Referencia de Bentos Costero de El Colegio de la Frontera Sur (ECOSUR).

The following abbreviations are used on the figures: A, Antenna; C, cephalon; Cx, Coxa; Dv, dorsal view; Fa, flagellar article 2 and 3; Gp, gnathopod; Ha, habitus; Ll, lower lip; Lv, lateral view; Md, mandible; Mo, molar; Mp, maxilliped; Mx, maxilla; P, pereopod; Pa, peduncular article, inner view; Pe, pereonite 1, dorsally; Pl, pleopod; Ul, upper lip; Us, urosome; Vp, ventral projection, article 2, antenna 2.

Systematics

Order Amphipoda Latreille, 1816

Suborder Corophiidea Leach, 1814

Family Ischyroceridae Stebbing, 1899

Tribe Siphonoecetini Just, 1983

Caribboecetes Just, 1983

Caribboecetes progreso sp. nov.Figures 1–3

Caribboecetes sp. Paz-Ríos & Ardisson, 2013: 152.

Holotype. Male (dissected and drawn), 4.4 mm, Yucalpeten beach, Progreso, Yucatan State, Mexico, 21°16’36’’N, 89°42’49’’W (approx. to 80 m from the shoreline), 1 May 2002, sandy bottom, 1.2 m depth, col. A. Vega, E. Campos, M.T. Herrera-Dorantes and P.-L. Ardisson, CYMX-1-CP.

Paratypes. 7 males and 1 gravid female, data as for holotype, ECOSUR0150. 3 males and 4 females, Chuburna beach, Progreso, Yucatan State, Mexico, 21°14’30’’N, 89°51’54’’W (approx. to 120 m from the shoreline), 18 September 2002, sandy bottom, 1.8 m depth, col. A. Vega, E. Campos, M.T. Herrera-Dorantes and P.-L. Ardisson, CYMX-2-CP. 1 male, Uaymitun beach, Progreso, Yucatan State, Mexico, 21°19’08’’N, 89°28’30’’W (approx. to 160 m from the shoreline), 3 May 2003, sandy bottom, 2 m depth, col. A. Vega, E. Campos, M.T. Herrera-Dorantes and P.-L. Ardisson, CYMX-3-CP. 1 male, northeastern continental shelf of Yucatan, Mexico, 21°47’44.4’’N, 89°30’1.5’’W, 9 April 2004, sandy bottom, 19.6 m depth, col. C.E. Paz-Ríos, CYMX-61-PY. 1 male, 1 female, northeastern continental shelf of Yucatan, Mexico, 21°42’42.3’’N,

Zootaxa 3652 (3) © 2013 Magnolia Press · 371A NEW SPECIES OF CARIBBOECETES FROM THE GULF OF MEXICO


88°00’39.9’’W, 9 April 2004, sandy bottom, 19.3 m depth, col. C.E. Paz-Ríos, CYMX-62-PY. 1 male (dissected), Xtul beach, Progreso, Yucatan State, Mexico, 21°15’17’’N, 89°49’39’’W, 9 January 2008, sandy bottom, 2 m depth, col. C.E. Paz-Ríos and M.T. Herrera-Dorantes, CYMX-4-CP. 2 male (dissected), Xtul beach, Progreso, Yucatan State, Mexico, 21°15’17’’N, 89°49’39’’W, 10 April 2008, sandy bottom, 2 m depth, col. C.E. Paz-Ríos and M.T. Herrera-Dorantes, CYMX-5-CP.

Type locality. Progreso, Yucatan State, Mexico.Etymology. The species name is derived from the type locality of the species.Diagnosis. Antenna 2 as long as cephalon and pereon combined; setae on peduncular articles 5 in a single row.

Eyelobes with short apical projection. Basis of gnathopod 2 with inflated triangular anterolateral flange. Anterior quarter of lateral surface on coxa plate 7 and basis of pereopod 7 densely setose.

Description. Based on male holotype (CYMX-1-CP) and three male paratypes (CYMX-4-CP; CYMX-5-CP). Rostrum acute, triangular, not reaching beyond eyelobes, straight though slightly depressed in lateral view. Base of rostrum microsetulose. Eyelobes slightly pedunculate, longer than broad, apex with pointed projection and a few setae (3–4). Cephalon and first 4–5 pereonites dorsally with 2–3 transverse rows of densely set polyflagellate setules.

Antennae 1 slightly shorter than antenna 2, article 1–2 extremely densely setose in male, less densely setose in female; sensory organs (aesthetascs) on last 8 articles. Antennae 2, ventral process of article 2 with 2–3 robust setae on rounded apex and lateral row of a few setae (5); peduncular article 3 medially with 1 distal, 1 midmedial, and 2–3 proximal straight robust setae; article 4 with dorsal setae in 10–12 short, transverse rows; article 5 with dorsolateral setae in dense longitudinal row contiguous with row of setae on flagellar articles 1–2; flagellar article 2 with 1 ventroapical robust seta and 2 on flagellar article 3.

Lower lip with simple and rounded inner lobes; rounded outer lobes with 4 medial robust setae. Upper lip with small apical notch. Mandible with triturative molar, well developed incisor, 5-toothed lacinia mobilis and 3 raker setae; palp bi-articulate with article 1 plumose (omitted setules in the drawing) and article 2 tiny. Maxilla 1 with inner plate reduced; outer plate with 7 apical bifurcate robust setae; palp with 4–5 apical robust setae and 1–2 setae. Maxilla 2 with subequal, rounded plates. Maxilliped with outer plate bearing 7 slender, oval, medial robust setae (middle ones with a few small coarse denticles) and 2–3 long plumose setae; inner plate with 3 oval, undentated medial robust setae and short plumose setae.

Coxal plate 1 triangular, longer than broad; plate 2 nearly square; plate 3–4 tapering, rounded apex, apical margin and most of anterior margin dentate, distolateral surface densely covered by acute cuticular scales; plates 5–6 with rounded, slightly upward turned posteroproximal lobe; plate 7 densely setose on anterolateral surface.

Gnathopod 1 simple; carpus with 1 posterodistal robust seta; propodus with 1 distal and 1 medial robust seta on posterior margin, medial surface with 2–3 transverse rows of short, stiff setae; dactylus with 4–5 posterior teeth. Gnathopod 2 simple; basis with inflated triangular anterolateral flange; propodus with posteroapical tooth, 5 medial robust setae along posterior margin (4 small and 1 large), and 1 distal robust seta on posterior margin; dactylus with 3–5 posterior teeth. Pereopod 3 and 4 pairwise similar; basis of pereopod 3 slightly wider and with much denser anterior and posterior setation than in pereopod 4; merus produced anterodorsally, anterodorsal lobe reaching midway along propodus anterior margin; robust setae row (5–7) on posterior margin of carpus; dactylus almost as long as combined length of carpus and propodus. Pereopod 5 and 6 pairwise identical; both pereopods geniculate at carpus; carpus with dense robust setae row (8–10) on posterior margin and apical and lateral surfaces covered by cuticular scales; dactylus short, curved and without accessory tooth. Pereopod 7 longest; basis oval with anterior and posterior margins densely fringed with long, heavily plumose setae and one quarter of anterolateral surface densely setose.

Pleopods 1–3 identical; peduncle broadly expanded medially with 2 serrated coupling spines.Uropod 1 biramous; peduncle with 4–7 dorsolateral setae and 4–6 distal robust setae; inner ramus 1/3 the

length of outer ramus with 5–7 lateral and apical robust setae; outer ramus 3 times as long as broad, with 14–17 robust setae along lateral and apical margin, medioapical robust seta notably longer than other robust setae. Uropod 2 absent. Uropod 3 without rami; peduncle apically rounded with numerous lateral seta of varying length and one apical robust seta.

Telson with oval field of spines covering ca. 1/2 of lateral margin, dorsal surface with a few setae proximal to normal setation.

PAZ-RÍOS & ARDISSON372 · Zootaxa 3652 (3) © 2013 Magnolia Press


FIGURE 1. Caribboecetes progreso sp. nov., holotype. Scale for Ha represent 1 mm, scales for Fa and Pe represent 0.1 mm, remainder represent 0.2 mm.



FIGURE 2. Caribboecetes progreso sp. nov., holotype. Scale represents 0.2 mm.



FIGURE 3. Caribboecetes progreso sp. nov., holotype, except for Us’: paratype (CYMX-4-CP). Scale for Cx3 represent 0.1 mm, remainder represent 0.2 mm.

Variation. There is some slight sexual dimorphism; female is similar to male, but with smaller size, antennae shorter and less setose and pereopods with fewer robust setae.

Remarks. A morphological comparison among the Caribboecetes species is presented in Table 1. Caribboecetes progreso sp. nov. is more similar to C. barbadensis, C. justi and especially to C. jenikarpae. The new species differs from C. barbadensis by the 2–3 transverse rows of setules dorsally on cephalon and first 4–5 pereonites, dorsolateral setae on peduncular and flagellar articles of antenna 2 in a single row, eyelobes slightly pedunculate, outer plate of maxilliped with oval medial robust setae, inflated triangular anterolateral flange on basis of gnathopod 2, coxal plate 3–4 with distal lateral surface covered with cuticular scales, and anterior lateral surface of coxal plate 7 and basis of pereopod 7 setose. It differs from C. justi by the rostrum not reaching eyelobes,



2–3 transverse rows of setules dorsally on cephalon and first 4–5 pereonites, dorsolateral setae on peduncular and flagellar articles of antenna 2 in a single row, eyelobes slightly pedunculate, outer lobes of lower lip with 4 medial robust setae, mandibular palp bi-articulate, outer plate of maxilliped with a few small coarse denticles on medial robust setae, coxal plate 3–4 with distal lateral surface covered with cuticular scales, and anterior lateral surface of coxal plate 7 and basis of pereopod 7 setose. It differs from C. jenikarpae by the outer lobes of lower lip with 4 medial robust setae, inflated triangular anterolateral flange on basis of gnathopod 2, and anterior lateral surface of coxal plate 7 and basis of pereopod 7 setose, which covers a half in C. jenikarpae and one quarter in C. progresosp. nov.

Distribution. Central coast of the Yucatan State, Mexico; northeastern continental shelf of the Yucatan Peninsula.

Ecological notes. Caribboecetes progreso sp. nov. has only been collected on sandy bottoms and it has been found in empty gastropods shell, mainly of the genus Olivella. It has a depth range from the intertidal zone to shallow continental shelf (19.6 m).

FIGURE 4. Abundance of Caribboecetes progreso sp. nov. in the type locality. A: First study, abundance by transect and sampling campaign; B: first study, abundance by distance from the shoreline and sampling campaign; C: third study, abundance by month.

The new species has been collected since the year 2002 on the central coast of the Yucatan State, Mexico, where it has achieved great abundance (Ardisson & Castillo-Fernández 2004). The spatial and temporal variation of abundance is displayed here for the first time (Figure 4); the total number of specimens in the first study was 49,992. Of this study it is observed the higher average abundance among transects was recorded during the sampling of April-May 2002, before the hurricane Isidore struck on September 2002 (Figure 4A); it is also



observed the higher average abundance was recorded at 60 m from shoreline, before the same hurricane. After the hurricane Isidore the abundance gradually increased among transects from the intertidal zone to shallow subtidal, with the higher value at 140–160 m from shoreline (Figure 4B). In the third study, the influence of the regional weather seasonality on population of C. progreso sp. nov. is observed; the total number of specimens in this study was 8,771. The higher average monthly abundance was recorded in April, a month characterized by a water column warmest and stable (proper to dry season); the lowest average abundance was recorded from August to February, a months characterized by a water column coolest and turbulent (proper to cold fronts season).

Geographical distribution of Caribboecetes

In order to cover the whole distribution of the genus and update the information, below is displayed the distribution for the six nominal species and two genera (Figure 5). Distribution is based on records from Just (1983, 1984), Ortiz and Lemaitre (1997), Díaz and Martín (2001), LeCroy (2007), Paz-Ríos and Ardisson (2013), and the present study.

FIGURE 5. Geographic distribution of the genus Caribboecetes.

There are two reports that were not included in the map of distribution because they are considered doubtful herein: the first one was a report of C. barbadensis in the Caribbean Sea of Venezuela by Ortiz et al. (2007); this one was an error during the typing; to date this species has not been found in that region (Y.J. Díaz, pers. comm.). The second one was a report of C. jenikarpae in the Gulf of California by García-Madrigal (2008); the occurrence of this species in that region was erroneously inferred from the cited source (Barnard & Karaman 1991) during compilation (M.S. García-Madrigal, pers. comm.).

There is also a possible record of C. jenikarpae in the eastern Pacific (Oaxaca, Mexico) by Peralta-García and García-Madrigal (2012), but it was not included in the present study because it has not been formally established on literature; the record was displayed on a meeting of carcinology.

Zoogeographically, the genus Caribboecetes displays an amphi-American distribution, with all species occurring in coastal marine waters (Just 1988). In the Atlantic it has a Caribbean affinity and in the Pacific a Tropical Eastern Pacific affinity. This pattern of distribution has been documented for other amphipod genera (e.g. Ampelisca, Neomegamphopus, Chelorchestia), identifying an eastern Pacific-Caribbean track of diversification,



which proposes an ancient connection between seas and a current isolation via emergence of the Central America isthmus as a major vicariant event (McKinney 1977; Foster et al. 2009; Myer & Lowry 2009; Neigel 2009).

Particularly, an outcome of that vicariant event was the final closure of the Isthmus of Panama 3.1 million years ago (Coates & Obando 1996), which resulted in the separation of related species pairs on the Atlantic and Pacific coast. Assuming that speciation of marine amphipods has been greatly due to their benthic life style and recruitment, their distribution patterns have been expected to depend on plate tectonics (Myers 1997; Myer & Lowry 2009). Thereby, the closure of the Isthmus of Panama as a biogeographic barrier for amphipods might be reflected by the separation of C. jenikarpae (Pacific coast) and C. progreso sp. nov. (Atlantic coast), two species of great similarity (e.g. the densely setose anterior lateral surfaces of coxal plate 7 and basis of pereopod 7).

Key to species of Caribboecetes (modified from Just 1984)

1 Basis of gnathopod 2 with strongly triangular upward anterolateral flange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2- Basis of gnathopod 2 without strongly triangular upward anterolateral flange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Rostrum not reaching beyond eyelobes; cephalon and first 4–5 pereonites dorsally with transverse rows of densely polyflagel-

late setules; coxal plate and basis of pereopod 7 with densely setose lateral surface . . . . . . . . . . . . . . . . . C. progreso sp. nov.- Rostrum reaching beyond eyelobes; cephalon and pereon with a few scattered simple setules; coxal plate 7 and basis of pereo-

pod 7 with smooth lateral surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. justi3 Cephalon and first 3–5 pereonites with transverse rows of setules dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4- Cephalon and pereon at most with scattered dorsal setules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Eyelobes with apical projection; peduncular article 5 and flagellar article 1 of antenna 2 with dorsolateral setae in a single dis-

tinct row; coxal plate and basis of pereopod 7 with densely setose lateral surface . . . . . . . . . . . . . . . . . . . . . . . . C. jenikarpae- Eyelobes apically rounded; dorsolateral setae on peduncular article 5 of antenna 2 not in a single row; coxal plate 7 and basis

of pereopod 7 with smooth lateral surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. barbadensis5 Rostrum curved strongly downward; article 1 of antenna 1 with broad, marginally setose lateral “wing” . . . . . . C. pterycornis- Rostrum straight; article 1 of antenna 1 normal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. intermedius

Acknowledgements

Thank you to Alfonso Ignacio Vega Moro and Eduardo Campos Bravo for their support on sampling survey, to María Teresa Herrera Dorantes (CINVESTAV, Mexico) for her support during the sampling and sorting of species, and to Delta Castillo Fernández for data processing in the first study. The first author thanks Jean Just (Zoological Museum, University of Copenhagen) for his valuable comments and suggestions on the manuscript. The sampling campaigns of the first study were supported by funds from CONABIO (project FB813/Y008/02) to P.L.A. The authors thank to anonymous reviewers who helped to improve the manuscript.

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caribboecetes progreso, a new species of sand-dwelling amphipod (amphipoda: corophiidea:...

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