Biodiversity and new records of benthic amphipods (Crustacea: Peracarida: Amphipoda) from coral reef protected natural areas in the Mexican Caribbean Sea

Biodiversidad y registros nuevos de los anfípodos bentónicos (Crustacea: Peracarida: Amphipoda) en áreas naturales protegidas de arrecifes de coral en el mar Caribe mexicano

Resumen

The Mesoamerican Caribbean Reef System (MCRS) is regarded as the second-largest reef system after Australia’s Great Barrier Reef and is one of 200 global priority ecoregions worldwide (Contreras-Silva et al., 2020). This huge ecosystem extends almost 1,600 km along the coastline of Mexico, Belize, Guatemala, and Honduras in the northwest sector of the Caribbean Sea and is comprised mainly of coral reefs along with mangroves, seagrass beds, and lagoon-inlet systems. The longest sector of this system is known as the Mexican Caribbean Sea (MEXCS), which is not only defined as a hotspot of endemic species, where numerous species can inhabit for feeding, nesting, and protection, but also has been impacted by coastal development, tourism, fishing (Perera-Valderrama et al., 2020), pollution, and invasive sargasso (Rodríguez-Martínez et al., 2020). Therefore, during the last 4 decades, the coastline ecosystems have been damaged by intensive urban and touristic expansion that has modified the structure, composition, and abundance of both benthic and pelagic communities.

As a result of natural ecological connectivity, knowledge of biodiversity in the MEXCS is required to design conservation, protection, and multidisciplinary management programs from those marine protected areas. So far, numerous organisms belonging to macroalgae, seagrass, mangroves, invertebrates, and chordates have been analyzed, either through species richness or abundance-distribution in this region (Ardisson et al., 2011; Contreras-Silva et al., 2020; Rioja-Nieto et al., 2019). In the case of peracarid crustaceans, which includes isopods, amphipods, tanaids, and 6 other orders, the taxonomic and ecological information is fragmentary throughout the MEXCS.

The order Amphipoda is comprised of almost 10,200 nominal species grouped in 6 suborders (Lowry & Myers, 2017; Horton et al., 2021). These peracarid crustaceans are associated with hard and soft bottoms, inhabiting areas from the coastline to the abyssal plain. Because of their abundance and wide-ranging geographic distribution, such benthic amphipods are involved in the trophic dynamics of marine communities as grazers, filter feeders, predators, scavengers, and prey (LeCroy et al., 2009). In coral reef ecosystems, amphipods are associated with sediments, coral rubble, seagrass beds, macroalgae mats, and sessile invertebrates such as corals, sponges, bryozoans, echinoderms, and tunicates, among others (Thomas, 1993a). 

Due to their ecological importance, numerical abundance, and sensitivity to a variety of toxins and pollutants, amphipod crustaceans are recognized as bioindicators in a broad variety of ecosystems, particularly in the tropics (Thomas, 1993b). Nevertheless, their incorporation into bioassessment programs on coral reefs and coastlines depends on taxonomic studies and inventories being completed (Monroy-Velazquez et al., 2017). 

During 2015 and 2016, 2 benthic sampling campaigns in the Coral Reef Puerto Morelos National Park (APM) and the Coral Reef Sian Ka’an Biosphere Reserve (SKAR), Quintana Roo, Mexico, were carried out to determine the biodiversity of benthic amphipods inhabiting several ecosystems and hard-soft bottoms. Also, the previously documented data on these peracarids from Contoy Island to the Xcalak Coral Reef were included in this paper. Therefore, the present study constitutes the first checklist of benthic amphipods along the MEXCS, including new records of benthic amphipod species in the Caribbean Sea, and provides a baseline of the existing marine amphipod biodiversity. 

Materials and methods

The MEXCS is located in the northwest Caribbean Sea, along the coastline of the Quintana Roo State, Mexico, spreading from Contoy Island on the northern side to Xcalak Coral Reef to the south (Fig. 1). It includes 16 protected marine areas (PMAs), which have been characterized as flora and fauna protection areas, national parks, biosphere reserves, and one sanctuary (Rioja-Nieto et al., 2019). According to Contreras-Silva et al. (2020), this Mexican sector of the Caribbean Sea is divided into 5 main regions, Northern, Center, Southern, Cozumel, and Banco Chinchorro. The PMA belongs to the Northern region, with a total area of 9,067 ha, 21 km in length, and 4.5 km seaward in addition to several well-preserved reef sites and extensive areas of seagrass beds (Monroy-Velázquez et al., 2017; Perera-Valderrama et al., 2020). Moreover, the Center region comprises both the Sian Ka’an Biosphere Reserve and the SKAR, which has a total marine area of 33,566 ha, 110 km length, and a maximum coral growth depth of 25 m. 

The information that the present study compiled on benthic amphipods arises essentially from the analysis of samples collected from macroalgae mats, seagrass beds, sponges, cnidarians, coral rubble, soft bottom, and wood debris from the APM and the SKAR during July 2015 and June 2016 (DGOPA.01024.110213.0236 and PPF/DGOPA-051/15 permits granted) as part of the “Macrocrustáceos asociados a los arrecifes de coral” project, which was supported by the Universidad Nacional Autónoma de México-DGAPA-PAPIIT. Such substrates were removed from 14 sample sites in both coral reefs by SCUBA diving between 0.5 and 25 m depth. The crustaceans associated with these substrates were separated, put into plastic bags, and preserved with 70% ethanol. Amphipods were divided from each crustacean sample and transported to the Facultad de Estudios Superiores Iztacala (FESI-UNAM). These specimens were examined and dissected using a stereoscopic microscope Motic SMZ-168 and optical microscope Leica DM750, both with camera lucida, at the Laboratorio de Crustáceos. 

The taxonomic analysis was essentially based on Barnard and Karaman (1991), LeCroy (2000, 2002, 2004, 2007), Lowry and Stoddart (1997), Ortiz et al. (2014), and Thomas and Klebba (2007). The classification system of Lowry and Myers (2017) was adopted for this study. The specimens collected were deposited in the Colección Nacional de Crustáceos (CNCR), Instituto de Biología, (UNAM-México). 

For the whole amphipod assemblage identified from both the APM and SKAR, specific amphipod richness and the total and relative abundances of each species were quantified, either for family, substrate, or site; additional information for each novel species recognized as a new record with geographic extension in the MEXCS, such as material examined, date, spatial distribution, depth, and substrate type, were provided. Also, new records of amphipods from the Caribbean Sea were documented to elucidate information regarding novel species, material examined, geo-referenced coordinates, the main morphological features, and previous geographic distribution where each species had been documented. 

Finally, in addition to the amphipod species collected from the AMP and SKAR, we present previous amphipod records to summarize the benthic amphipods checklist which we compiled using published references from the MEXCS and verified according to the comprehensive taxonomic list for the ecoregions of the Caribbean Sea published by Ortiz et al. (2007) and Martin et al. (2013). Every suborder, family, genus, and species are presented in an alphabetical rather than phyletic order, and the references where every species was reported along the MEXCS are included in the checklist. 

Results

The present study documented a total of 120 nominate species grouped into 70 genera and 34 families, belonging to Amphilochidea (14 families), Colomastigidea (1 family), and Senticaudata (19 families) suborders. The most diversified amphipod families in this large sector of the Caribbean Sea were (in decreasing order): Maeridae, Aoridae, Leucothoidae, Ampeliscidae, Ampithoidae, Amphilochidae, Melitidae, Lysianassidae, and Pontogeneiidae. Also, and as a result of the 2 oceanographic campaigns carried out in APM and SKAR, Quintana Roo, not only 82 benthic amphipod species were identified, out of which 41 species had not been previously recognized along the MEXCS, but also 5 species represented new amphipod records from the Caribbean Sea. 

New records of amphipods from the Caribbean Sea

Shoemakerella lowryi Gable & Lazo-Wasem, 1990. Material examined: APM; 20°50’28.57” N, 86°52’25.45” W; 12 m depth; 6-VI-2015; 8 specimens, M. Lozano-Aburto and I. Winfield, colls.; I. Winfield and M. Ortiz det., CNCR36044. This species can be recognized by the dorsal process on the peduncle of uropod 3, the proportion between basis and ischium of gnathopod 2, and the shape of basis of pereopod 7. Also, S. lowryi differs from S. cubensis in the relative lengths of propodus of peraeopod 7 (length 9.5 x width vs. length 5-6 x width in S. cubensis). This shallow species had been only documented in Bermuda (Gable & Lazo-Wasem, 1990; WoRMS, 2021), the occurrence of this species in APM associated with coral rubble, represents the first new record from the Caribbean Sea.

Stenothoe minuta Holmes, 1905. Material examined: SKAR; 20°04’09.30” N, 87°28’09.30” W; 13 m depth; 19-VI-2016; 2 specimens, M. Lozano-Aburto and I. Winfield, colls., I. Winfield and M. Ortiz det., CNCR36050. This stenothoid species is identified by outer ramus of uropod 2 shorter than inner ramus, gnathopod 1 propodus posterior margin straight and shorter than palmar margin, gnathopod 2 palmar angle weakly angled-unrounded, without spinose lobe, peraeopod 7 basis broad with posterior margin convex, uropod 3 peduncle longer than article 1 of ramus. This shallow species had been only documented in several shoreline ecosystems along the Gulf of Mexico, from northeast Florida to the southwest of the Gulf of Mexico (LeCroy et al., 2009). The presence of S. minuta in macroalgae mats in SKAR is the second new record from the Caribbean Sea.

Bemlos tigrinus (A.A. Myers, 1979). Material examined: APM; 20°52’06.9” N, 86°50’51.6” W; 10 m depth; 6-VI-2015; 5 specimens, M. Lozano-Aburto and I. Winfield, colls., I. Winfield and M. Ortiz det., CNCR36068. This species can be distinguished by peraeon segments 2-4 with mid-ventral sternal process, third the largest, pigmentation pattern on body composed of dorsal dark bands combined along the body, gnathopod 2 merus and carpus setose on anterior margin, peraeopods 5-6 basis with margin short simple setae. This shallow species had been recorded from West Florida Bay, NW Gulf of Mexico (LeCroy et al., 2009), a consequence of such species, which was collected from APM in T. testudinum, is a new record for the Caribbean Sea.

Protohyale (Protohyale) frequens Bousfield & Hendrycks, 2002. Material examined: APM; 20°52’23.43” N, 86°51’9.95” W; 10 m depth; 9-VI-2015; 140 specimens, M. Lozano-Aburto and I. Winfield, colls., I. Winfield and M. Ortiz det., CNCR36082; SKAR; 20°00′40.07″ N, 87°28′07.41″ W; 3 m depth; 18-VI-2016; 5 specimens, M. Lozano-Aburto and I. Winfield, colls., I. Winfield and M. Ortiz det., CNCR36083. This species can be distinguished by the antenna 2 flagellum long, gnathopod 1 propodus elongate and subrectangular, gnathopod 2 propodus with anterior margin lacking robust setae, uropod 1 peduncular distolateral spine short, uropod 3 ramus slightly shorter than peduncle. This shallow species had only been recorded from coastline habitats, NE Gulf of Mexico (LeCroy, 2000), therefore, the finding of this species in APM, and SKAR, in wood debris, wood dock, and macroalgae mats, represents a new record from the Caribbean Sea. 

Melita elongata Sheridan, 1980. Material examined: SKAR; 20°00’40.07” N, 87°28’07.41” W; 4 m depth; 18-VI-2016; 2 specimens, M. Lozano-Aburto and I. Winfield, colls., I. Winfield and M. Ortiz det., CNCR36104. This species can be distinguished by gnathopod 1 parachelate, gnathopod 2 oval, female with anterior lobe of coxa 6 with a large hook backward, telson without robust dorsal setae. According to LeCroy et al. (2009), it is distributed along the coastline of the Gulf of Mexico, from W Florida to Términos Lagoon. This species was found in coral rubble and soft bottoms in SKAR, consequently, it is a new record from the Caribbean Sea.

Benthic amphipods from coral reefs APM and SKAR

All amphipod species collected from both coral reefs’ protected areas were organized into 2 suborders, 27 families, 44 genera, and 82 nominal species. Particularly, APM included 58 species with 2,469 specimens, and 53 species with 1,319 organisms in SKAR, out of which 27 amphipod species were distributed in both protected areas, also Maeridae, Aoridae, and Leucothoidae were the most specious families (Table 1). On the other hand, 80% of the total abundance were accumulated for Chevaliidae (19%), Maeridae (16%), Ampithoidae (15%), Leucothoidae (14%), Pontogeneiidae (9%), and Ischyroceridae (8%) from APM, and Maeridae (33%), Ampithoidae (17%), Chevalidae (14%), Photidae (12%), and Aoridae (9%), in SKAR (Table 1).

The dominant species of benthic amphipod in AMP, which represented almost 75% of abundance, were Chevalia mexicana, Leucothoe saron, Ampithoe longimana, A. marcuzzii, Ericthonius brasiliensis, Nasageneia yucatanensis, Elasmopus balkomanus, E. pocillimanus, and Protohyale frequens; whereas in SKAR, Quadrimaera miranda, Q. quadrimana, C. mexicana, Latigammaropsis atlantica, Elasmopus rapax, A. longimana, Leucothoe barana, L. kensleyi, L. ashleyae, L. garifunae, Concarnes concavus, and Hourstonius laguna, which accumulated 80% of such relative abundance. 

The abundance and biodiversity of the whole amphipod assemblage in AMP were predominantly associated with macroalgae mats and coral rubble, each one with 41%, reducing in sponges (13%), octocoral (12%), seagrass beds (3%), and soft bottoms (> 1%). In contrast, the substrate prevalence in SAKR included the macroalgae mats with 75%, and 25% was distributed among coral rubble, octocoral, soft bottoms, seagrass beds, and sponges.

Table 1

Families, species, and total abundance of each benthic amphipods. AMP, coral reef Puerto Morelos; SKAR, coral reef Sian Ka’an. Suborders and voucher numbers (CNCR) were included. 

Suborder/Family	Species				CNCR	APM	SKAR
Amphilochidea
Ampeliscidae	Ampelisca bicarinata Goeke & Heard, 1983	36010	4
	Ampelisca burkei J.L. Barnard & Thomas, 1989	36011 36012	12	3
	Ampelisca schellenbergi Shoemaker, 1933	36013		4
Amphilochidae	Apolochus casahoya (Mckinney, 1978)	36014	2
	Apolochus delacaya (Mckinney, 1978)	36015	4
	Apolochus pillaii (Barnard & Thomas, 1983)	36016	6
	Gitana dominica Thomas and Barnard, 1990	36017 36018	2	3
	Hourstonius laguna (McKinney, 1978)	36019 36020	3	4

Table 1. Continued
Suborder/Family	Species		CNCR	APM	SKAR
	Hourstonius tortugae (Shoemaker, 1942)	36021	29
Bateidae	Batea campi (Ortiz, 1991)		36022	12
	Batea carinata (Shoemaker, 1926)		36023 36024	4	2
	Batea catharinensis Muller, 1865		36025	15
Cyproideidae	Hoplopheonoides obesa Shoemaker, 1956	36026	3
Leucothoidae	Anamixis cavatura Thomas, 1997		36027		2
	Anamixis hanseni Sttebing, 1897		36028		2
	Leucothoe ashleyae Thomas & klebba, 2006	36029 36030	59	5
	Leucothoe barana Thomas & Klebba, 2007	36031 36032	30	17
	Leucothoe garifunae Thomas & Klebba, 2007	36033 36034	8	3
	Leucothoe kensleyi Thomas & Klebba, 2005	36035 36036	4	6
	Leucothoe laurensi Thomas & Ortiz, 1995	36037		3
	Leucothoe saron Thomas & Klebba, 2007	36038	244
	Leucothoe spinicarpa (Abildgaard, 1789)	36039		15
	Leucothoe ubouhu Thomas & Klebba, 2007	36040		2
Liljeborgiidae	Liljeborgia bousfieldi McKinney, 1979	36041	3
Lysianassidae	Concarnes concavus Shoemaker, 1933	36042		10
	Lysianopsis alba (Holmes, 1905)		36043		2
	Shoemakerella lowryi Gable & Lazo-Wasem, 1990	36044	8
	Tantena zlatarskii Ortiz, Lalana & Varela, 2007	36045		2
Oedicerotidae	Westwoodilla longimana Shoemaker, 1934	36046		2
Phoxocephalidae	Eobrolgus spinosus (Holmes, 1905)		36047	3
	Metharpinia floridana (Shoemaker, 1933)	36048	5
Stenothoidae	Stenothoe gallensis Walker, 1904		36049	3
	Stenothoe minuta Holmes, 1905		36050		2
	Stenothoe valida Dana, 1852		36051		2
Synopiidae	Synopia ultramarina Dana, 1853		36052		2
Senticaudata
Ampithoidae	Ampithoe longimana Smith, 1873		36053 36054	225	122
	Ampithoe marcuzzii Ruffo, 1954		36055	125
	Ampithoe ramondi Audouin, 1826		36056 36057	17	3
	Ampithoe valida Smith, 1873		36058	10
	Pseudamphithoides bacescui Ortiz, 1976	36059		102
Aoridae	Bemlos dentischium (A.A. Myers, 1977)	36060		18
	Bemlos inermis Myers, 1979		36061 36062	2	6
	Bemlos kunkelae (A.A. Myers, 1977)	36120		4
	Bemlos longicornis (A.A. Myers, 1989)	36063	18
	Bemlos mackinneyi (A.A. Myers, 1978)	36064		2
	Bemlos spinicarpus (Pearse, 1912)		36065 36066	11	6
	Bemlos tigrinus (A.A. Myers, 1979)		36067	5
	Bemlos unicornis (Bynum & Fox, 1977)	36068 36069	11	4
	Globosolembos smithi (Holmes, 1905)	36070		4
	Lembos unifasciatus Myers, 1977		36071 36072	14	63
	Plesiolembos rectangulatus (A.A. Myers, 1977)	36073 36074	4	6
Chevaliidae	Chevalia carpenteri Barnard & Thomas, 1987	36075		2
	Chevalia mexicana Pearse, 1913		36076 36077	477	188
Hadziidae	Dulzura schoenerae (Fox, 1973)		36078	3
Hyalidae	Apohyale media (Dana, 1853)		36079		5
	Parhyale hawaiensis (Dana, 1853)		36080		4
	Protohyale (Protohyale) frequens Bousfield & Hendrycks, 2002	36081 36082	140	5
Ischyroceridae	Ericthonius brasiliensis (Dana, 1853)	36083 36084	204	53
	Ericthonius rubricornis (Stimpson, 1853)	36085	2
Maeridae	Ceradocus (Denticeradocus) sheardi Shoemaker, 1948		36086	3
	Ceradocus shoemakeri Fox, 1973		36087		2
	Elasmopus balkomanus Tomas & Barnard, 1988	36088 36089	170	4
	Elasmopus levis (S.I. Smith in Verrill, 1873)	36090 36091	13	4
	Elasmopus pocillimanus (Spence Bate, 1862)	36092	146
	Elasmopus rapax Costa, 1853		36093 36094	7	7
	Elasmopus thomasi Ortiz & Lalana, 1994	36095	3
	Quadrimaera inaequipes (A. Costa in Hope, 1851)	36096	3
	Quadrimaera miranda (Ruffo, Krapp-Schickel & Gable, 2000)	36097 36098	14	196
	Quadrimaera pacifica (Schellenberg, 1983)	36099	10
	Quadrimaera quadrimana (Dana, 1852)	36100 36101	17	226
	Spathiopus looensis Thomas & Barnard, 1985	36102	3
Melitidae	Melita elongata Sheridan, 1980		36103		2
Neomegamphopidae	Neomegamphopus hiatus Barnard & Thomas, 1987	36104	3
Nuuanuidae	Nuuanu muelleri Ortiz, 1976		36105 36106	2	4
Phliantidae	Pariphinotus seclusus (Shoemaker, 1933)	36107	73
Photidae	Gammaropsis chelifera (Chevreux, 1901)	36108 36109	4	30
	Latigammaropsis atlantica (Stebbing, 1888)	36110 36111	72	161
Podoceridae	Podocerus kleidus Thomas & Barnard, 1992	36112 36113	2	7
Pontogeneiidae	Nasageneia bacescui Ortiz & Lalana, 1994	36114 36115	10	4
	Nasageneia yucatanensis Ledoyer, 1986	36116 36117	195	10
	Tethygeneia longleyi (Shoemaker, 1933)	36118	10
Talitridae	Chelorchestia forceps Smith & Heard, 2001	36119		2

New records and geographical extension along the MEXCS 

1. Ampelisca burkei Barnard & Thomas, 1989. Material examined: APM; 15 m depth; 6-VI-2015; 12 specimens; SKAR; 8 m depth; 16-VI-2016; 3 specimens. This shallow species inhabits macroalgae mats in APM and SKAR associated with coral rubble. 

2. Apolochus delacaya (Mckinney, 1978). Material examined: APM; 9 m depth; 4-VI-2015; 4 specimens. This shallow species occurs in the APM in coral rubble and macroalgae mats. 

3. Apolochus pillaii (Barnard & Thomas, 1983). Material examined: APM; 15 m depth; 7-VI-2015; 6 specimens. This shallow species occurs in the APM in coral rubble and macroalgae mats. 

4. Hourstonius laguna (McKinney, 1978). Material examined: APM; 8 m depth; 5-VI-2015; 3 specimens; SKAR; 12 m depth; 16-VI-2016; 4 specimens. This shallow species occurs associated with macroalgae mats and coral rubble. 

5. Batea campi (Ortiz, 1991). Material examined: APM; 8 m depth; 6-VI-2015; 12 specimens. This shallow species is recorded from APM in coral rubble and macroalgae mats. 

6. Batea catharinensis Muller, 1865. Material examined: APM; 12 m depth; 4-VI-2015; 15 specimens. This shallow species occurs in the APM associated with coral rubble. 

7. Hoplopheonoides obesa Shoemaker, 1956. Material examined: APM; 12 m depth; 4-VI-2015; 3 specimens. This shallow species is restricted to APM associated with coral rubble. 

8. Anamixis hanseni Sttebing, 1897. Material examined: SKAR; 12 m depth; 17-VI-2016; 2 specimens. This shallow species occurs from macroalgae mats in SKAR. 

9. Leucothoe ashleyae Thomas & Klebba, 2006. Material examined: APM; 15 m depth; 6-VI-2015; 59 specimens; SKAR; 8 m depth; 15-VI-2016; 5 specimens. This shallow species inhabits both APM and SKAR in macroalgae mats. 

10. Leucothoe barana Thomas & Klebba, 2007. Material examined: APM; 20°50’28.57” N, 86°52’25.45” W; 12 m depth; 6-VI-2015; 30 specimens; SKAR; 12 m depth; 18-VI-2016; 17 specimens. This shallow species is found in APM and SKAR associated with coral rubble, macroalgae mats, and T. testudinum. 

11. Leucothoe garifunae Thomas & Klebba, 2007. Material examined: APM; 10 m depth; 6-VI-2015; 8 specimens; SKAR; 11 m depth; 17-VI-2016; 3 specimens. This shallow species is found from both APM and SKAR associated with wooden piles and macroalgae mats. 

12. Leucothoe kensleyi Thomas & Klebba, 2005. Material examined: APM; 10 m depth; 6-VI-2015; 4 specimens; SKAR; 9 m depth; 16-VI-2016; 6 specimens. This shallow species is found from both APM and SKAR associated with T. testudinum. 

13. Leucothoe saron Thomas & Klebba, 2007. Material examined: APM; 8 m depth; 6-VI-2015; 244 specimens. This shallow species is found in APM in sponges Callyspongia aculeata, Ircinia strobilina, and coral rubble. 

14. Leucothoe ubouhu Thomas & Klebba, 2007. Material examined: SKAR; 13 m depth; 17-VI-2016; 2 specimens. This shallow species is restricted to SKAR in macroalgae mats. 

15. Tantena zlatarskii Ortiz, Lalana & Varela, 2007. Material examined: SKAR; 10 m depth; 18-VI-2016; 2 specimens. This shallow species is restricted to SKAR in soft bottoms. 

16. Westwoodilla longimana Shoemaker, 1934. Material examined: SKAR; 11 m depth; 18-VI-2016; 2 specimens. This shallow species is restricted to SKAR in macroalgae mats. 

17. Eobrolgus spinosus (Holmes, 1905). Material examined: APM; 10 m depth; 8-VI-2015; 3 specimens. This shallow species is restricted to APM associated with coral rubble and seagrass beds. 

18. Stenothoe gallensis Walker, 1904. Material examined: APM; 7 m depth; 6-VI-2015; 3 specimens. This shallow species is restricted to APM associated with macroalgae mats. 

19. Stenothoe valida Dana, 1852. Material examined: SKAR; 15 m depth; 18-VI-2016; 2 specimens. This shallow species is documented from SKAR associated with macroalgae mats. 

20. Ampithoe marcuzzii Ruffo, 1954. Material examined: APM; 11 m depth; 7-VI-2015; 125 specimens. This shallow species is found in APM in coral rubble and sponge Ircinia felix. 

21. Ampithoe valida Smith, 1873. Material examined: APM; 12 m depth; 8-VI-2015; 10 specimens. This shallow species is recorded from APM in coral rubble. 

22. Pseudamphithoides bacescui Ortiz, 1976. Material examined: SKAR; 16 m depth; 16-VI-2016; 102 specimens. This shallow species is recorded from SKAR in macroalgae mats. 

23. Bemlos dentischium (A.A. Myers, 1977). Material examined: SKAR; 9 m depth; 17-VI-2016; 18 specimens. This shallow species is found in SKAR associated with coral rubble and macroalgae. 

24. Bemlos inermis Myers, 1979. Material examined: APM; 8 m depth; 5-VI-2015; 2 specimens; SKAR; 10 m depth; 17-VI-2016; 6 specimens. This shallow species is found in APM, and SKAR inhabits coral rubble, macroalgae mats, and T. testudinum.

25. Bemlos kunkelae (A.A. Myers, 1977). Material examined: SKAR; 12 m depth; 17-VI-2016; 4 specimens. This shallow species is recorded from SKAR associated with coral rubble, and macroalgae mats.

26. Bemlos longicornis (A.A. Myers, 1989). Material examined; APM; 12 m depth; 8-VI-2015; 18 specimens. This shallow species is found in APM inhabits in T. testudinum.

27. Bemlos mackinneyi (A.A. Myers, 1978). Material examined: SKAR; 12 m depth; 18-VI-2016; 2 specimens. This shallow species occurs in SKAR inhabits macroalgae mats.

28. Plesiolembos rectangulatus (A.A. Myers, 1977). Material examined: APM; 19 m depth; 8-VI-2015; 4 specimens; SKAR; 7 m depth; 16-VI-2016; 6 specimens. This shallow species had been collected in APM and SKAR associated with macroalgae mats and coral rubble.

29. Dulzura schoenerae (Fox, 1973). Material examined: APM; 15 m depth; 9-VI-2015; 3 specimens. So far, this shallow species is only recorded from APM associated with coral rubble.

30. Apohyale media (Dana, 1853). Material examined: SKAR; 4 m depth; 18-VI-2016; 5 specimens. This shallow species is found in SKAR inhabits macroalgae mats. 

31. Parhyale hawaiensis (Dana, 1853). Material examined: SKAR; 3 m depth; 18-VI-2016; 4 specimens. This shallow species is found in SKAR in macroalgae mats.

32. Elasmopus pocillimanus (Spence Bate, 1862). Material examined: APM; 10 m depth; 7- VI-2015; 146 specimens. So far, this shallow species is recorded in APM in macroalgae mats.

33. Elasmopus thomasi Ortiz & Lalana, 1994. Material examined: APM; 15 m depth; 8-VI-2015; 3 specimens. So far, this shallow species is found in APM associated with coral rubble.

34. Quadrimaera pacifica (Schellenberg, 1983). Material examined: APM; 10 m depth; 8-VI-2015; 10 specimens. So far, this shallow species is recorded from APM associated with T. testudinum.

35. Neomegamphopus hiatus Barnard & Thomas, 1987. Material examined: APM; 19 m depth; 9-VI-2015; 3 specimens. So far, this shallow species is recorded in APM associated with macroalgae mats. 

36. Gammaropsis chelifera (Chevreux, 1901). Material examined: APM; 7 m depth; 6-VI-2015; 4 specimens; SKAR; 5 m depth; 18-VI-2016; 5 specimens. It is found in APM, and SKAR associated with coral rubble, macroalgae mats, and soft bottoms. 

37. Podocerus kleidus Thomas & Barnard, 1992. Material examined: APM; 14 m depth; 9-VI-2015; 2 specimens; SKAR; 10 m depth; 18-VI-2016; 7 specimens. This shallow species is found in APM, and SKAR associated with macroalgae mats, and seagrass beds. 

38. Nasageneia bacescui Ortiz & Lalana, 1994. Material examined: APM; 9 m depth; 9-VI-2015; 10 specimens; SKAR; 4 m depth; 18-VI-2016; 4 specimens. This shallow species is reported in APM, and SKAR inhabits in macroalgae mats and coral rubble. 

39. Nasageneia yucatanensis Ledoyer, 1986. Material examined: APM; 10 m depth; 8-VI-2015; 195 specimens; SKAR; 15 m depth; 16-VI-2016; 10 specimens. This shallow species is reported in APM, and SKAR inhabits macroalgae mats and coral rubble.

40. Tethygeneia longleyi (Shoemaker, 1933). Material examined: APM; 10 m depth; 9-VI-2015; 10 specimens. So far, this shallow species is documented from APM associated with macroalgae mats and soft bottoms.

41. Chelorchestia forceps Smith & Heard, 2001. Material examined: SKAR; 3 m depth; 18-VI-2016; 2 specimens. This shallow species is found in SKAR associated with coral rubble.

Discussion

The present study contributes to the first inventory of benthic amphipod species along the MEXCS documented from 1977 to date, including those species collected and analyzed in this work from both the APM and the SKAR, Quintana Roo, in 2015 and 2016. Previously, Paz-Ríos et al. (2021) documented some species richness ranges in benthic amphipods for the Western Caribbean ecoregion. 

The MEXCS coastline includes beaches, rocky shores, seagrass beds, coral reefs, mangroves, and karstic bottoms, where the freshwater aquifer and seawater are frequently mixed (Rodríguez-Martínez et al., 2020). Such ecosystems make up a complex semi-continuous coral reef barrier with the key geomorphological features of back-reef, reef-crest, and front-reef (Rioja-Nieto & Álvarez-Filip, 2019). Throughout the MEXCS the benthic amphipods occur in diverse types of substrates, but are predominantly associated with coral rubble (88 species) and macroalgae mats (60 species), decreasing in species richness in seagrass beds and soft bottoms (22 species in each). According to the information summarized in this amphipod inventory, both the APM (38%, 83 species) and the SKAR (24%, 52 species), located in the north and central sectors of the MEXCS, represented the protected natural areas with the largest amounts and percentages of benthic amphipod species, with decreases in peracarids from the north sector to the south sector: Isla Mujeres (3%, 6 species), Isla Cozumel (7%, 17 species), Bahía Ascención (13%, 29 species), Banco Chinchorro (12%, 26 species), and Chetumal-Belice border (3%, 8 species). Besides, 84% of amphipod records belong to shallow water substrates (either soft-hard bottoms or living substrata), and 16% are from deep coral rubble (deeper than 25 m).

Although most coral reef works have documented tens of sponges hosting amphipod species belonging to Colomstigidae, Leucothoidae, Aoridae, and Maeridae, only Leucothoe spinicarpa, Leucothoe saron, Ampithoe marcuzzii, and Ericthonius brasiliensis have been recorded in sponges throughout the MEXCS. This fact is a result of the multidisciplinary management strategies proposed by the Protected Natural Areas Agency-Mexico, where the conservation and protection of numerous benthic invertebrates, such as sponges, represents a key component as a refuge for feeding, protection, and reproduction for several invertebrate species. Thus, knowledge of biodiversity and interactions between amphipod-sponges are fragmentary in the MEXCS.

Until now, 13 benthic amphipod species have been reported in most coral reef protected areas belonging to the MEXCS, out of which only Leucothoe spinicarpa, Lysianopsis alba, Elasmopus rapax, and E. levis occur throughout the whole MEXCS (Table 2). Recent analyses on biogeographic aspects showed 2 of these species (E. rapax and L. spinicarpa) are widely distributed across the warm Northwest Atlantic ecoregions (Paz-Ríos et al., 2021). Until the amphipod inventory is completed for the Mexican Caribbean Sea protected areas, we cannot determine if the geographic distribution pattern of such benthic crustaceans is continuous or if it is defined as “patches” in each coral reef protected area. Thomas (1993a) suggested that the ecological features of these amphipod assemblages in each coral reef ecosystem, such as substrate specificity and biological adaptations, should be analyzed by microhabitats rather than a huge macrohabitat, which is a hypothesis to be tested in future research.

Table 2

Checklist of benthic amphipods throughout MEXCS. IM, Isla Mujeres; APM, Arrecife Puerto Morelos; IC, Isla Cozumel; SKA, Arrecife Sian Ka’an; BA, Bahía Ascención; RBC, Reserva Biosfera Chinchorro; Ch-B, Chetumal-Belice; REF, referencia. 1, Monroy-Velázquez et al. (2017); 2, Monroy-Velázquez et al. (2020); 3, Escobar-Briones and Winfield (2003); 4, Oliva-Rivera (2003); 5, Oliva-Rivera (1998); 6, McKinney (1977); 7, Oliva-Rivera and Jiménez-Cueto (1992); 8, Cházaro-Olvera et al. (2021); 9, Oliva-Rivera and Jiménez-Cueto (1992); 10, LeCroy et al. (2009); 11, Wildish and LeCroy (2014). 

Taxon				IM	APM	IC	SKA	BA	RBC	Ch-B	REF
Ampeliscidae
Ampelisca abdita		*						1, 2
Ampelisca agassizi		*						1
Ampelisca bicarinata		*	*					1, 3
Ampelisca burkei		*		*				PS
Ampelisca holmesi			*					3
Ampelisca schellenbergi		*		*				1
Ampelisca vadorum			*					3
Ampelisca verilli			*					3
Amphilochidae
Apolochus casahoya			*				*	*	4, 5
Apolochus delacaya			*						PS
Apolochus neapolitanus			*		*	*	*		4, 6, 7
Apolochus pillaii		*						PS
Gitana dominica				*	*				3
Hourstonius laguna					*				PS
Hourstonius tortugae		*						1, 2
Atylidae
Nototropis minikoi	*			*				8
Bateidae
Batea campi		*						PS
Batea carinata		*		*	*	*		4
Batea catharinensis		*						PS
Batea cuspidata		*						1, 2
Cyproideidae
Hoplopheonoides obesa		*						PS
Leucothoidae
Anamixis cavatura		*		*				1, 2
Anamixis hanseni				*				PS
Leucothoe ashleyae		*						PS
Leucothoe barana			*		*				PS
Leucothoe garifunae		*		*				PS
Leucothoe kensleyi			*		*				PS
Leucothoe laurensi			*		*				1
Leucothoe saron			*						PS
Leucothoe spinicarpa		*	*	*		*	*		1, 2, 3, 5, 6, 7

Table 2. Continued
Taxon		IM	APM	IC	SKA	BA	RBC	Ch-B	REF
Leucothoe ubouhu					*				PS
Liljeborgiidae
Liljeborgia bousfieldi			*			*			1, 5
Lysianassidae
Concarnes concavus			*	*	*				1, 3
Dissiminassa dissimilis					*			5
Lysianopsis alba		*	*	*	*	*		1, 2, 3, 4, 5, 6, 7
Shoemakerella lowryi			*						PS
Tantena zlatarskii					*				PS
Ochlesidae
Curidia debrogania		*				*		1, 6
Idunella quintana		*					*		5, 9
Oedicerotidae
Perioculodes cerasinus		*						1, 2
Westwoodilla longimana				*				PS
Phoxocephalidae
Eobrolgus spinosus		*						PS
Metharpinia floridana		*						1, 2
Sebidae
Seba tropica			*					*	1
Stenothoidae
Stenothoe gallensis			*						PS
Stenothoe minuta				*				PS
Stenothoe valida				*				PS
Synopiidae
Synopia ultramarina			*		*	*	*		1, 6, 3
Trion triocellatus			*						1
Colomastigidae
Colomastix janiceae		*						1
Colomastix pusilla					*			5
Ampithoidae
Ampithoe longimana	*	*		*				8
Ampithoe marcuzzii		*						PS
Ampithoe ramondi		*		*				1, 2
Ampithoe valida		*						PS
Cymadusa filosa					*	*		6, 7, 9
Pseudamphithoides bacescui				*				PS
Pseudampithoides incurvaria			*						1, 2
Aoridae
Bemlos dentischium					*				PS
Bemlos inermis		*		*				PS
Bemlos kunkelae				*				PS
Bemlos longicornis		*						PS
Bemlos mackinneyi				*				PS
Bemlos spinicarpus		*	*	*		*		1, 5
Bemlos tigrinus		*						PS
Bemlos unicornis		*		*				1
Bemlos waipio					*	*		6, 7
Globosolembos smithi		*		*				1, 2
Grandidierella bonnieroides			*		*	*	*	3, 5, 6, 7
Lembos unifasciatus		*		*				1, 2
Plesiolembos rectangulatus		*		*				PS
Chevaliidae
Chevalia aviculae		*	*					1, 2, 3
Chevalia carpenteri				*			*	10
Chevalia mexicana		*		*			*	10
Eriopisidae
Netamelita barnardi		*						1, 2
Gammaridae
Gammarus mucronatus		*						1
Hadziidae
Dulzura schoenerae		*						PS
Hyalidae
Apohyale media				*				PS
Parhyale hawaiensis				*				PS
Protohyale (Protohyale) frequens		*		*				PS
Ischyroceridae
Ericthonius brasiliensis		*		*	*	*		1, 5, 7, 9
Ericthonius rubricornis		*	*					3
Maeridae
Anamaera hixoni		*						1
Ceradocus (Denticeradocus) sheardi		*		*	*	*		1, 2, 5, 7
Ceradocus shoemakeri		*		*				1
Dulichiella appendiculata			*			*		3, 5
Dumosus atari							*	10
Elasmopus balkomanus		*		*				1
Elasmopus levis	*	*		*	*	*		1, 2, 5, 6, 7
Elasmopus pocillimanus		*						PS
Elasmopus rapax	*	*	*	*	*	*		1, 2, 3, 5, 6, 7
Elasmopus thomasi		*						PS
Maera jerrica		*	*			*		1, 3
Quadrimaera inaequipes		*			*			5
Quadrimaera miranda		*		*				1
Quadrimaera pacifica		*						PS
Quadrimaera quadrimana		*		*	*	*		2, 6, 9
Spathiopus looensis		*						1
Megaluropidae
Gibberosus myersi		*						1
Melitidae
Dulichiella appendiculata		*	*		*	*		3, 6, 9
Elasmopus fusimanus					*			6
Elasmopus levis					*	*		5
Elasmopus rapax			*		*	*		3, 5
Melita elongata		*						PS
Melita nitida					*	*		5, 9
Neomegamphopidae
Neomegamphopus hiatus		*						PS
Nuuanuidae
Nuuanu copillius					*			10
Nuuanu muelleri		*		*				2
Phliantidae
Pariphinotus seclusus		*			*	*	*	1, 2, 6
Photidae
Gammaropsis chelifera		*		*				PS
Latigammaropsis atlantica		*		*				1, 2
Podoceridae
Podocerus kleidus		*		*				PS
Pontogeneiidae
Nasageneia bacescui		*		*				PS
Nasageneia yucatanensis		*						PS
Pontogeneia barstchi					*	*		5, 6
Tethygeneia longleyi		*						PS
Talitridae
Chelorchestia forceps				*				PS
Mexorchestia carpenteri					*		*	11
Tethorchestia antillensis					*	*		5
Unciolidae
Rudilemboides naglei			*	*				3

Acknowledgments

To the PAPIIT program, sponsored by the Dirección General de Asuntos del Personal Académico, Universidad Nacional Autónoma de México, for the financial support. In addition, thanks are due to Comisión Nacional de Acuacultura y Pesca (Conapesca-DGOPA) for the collecting license (DGOPA.01024.110213.0236 and PPF/DGOPA-051/15). Additionally, we would like to thank to M. A. Lozano-Aburto for his support during the sampling and sorting of species.

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