Research Article |
Kristine N. White‡
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Corresponding author: Kristine N. White ( kristine.white@gcsu.edu ) Academic editor: Alan Myers
© 2024 Kristine N. White.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
White KN (2024) Caribbean Amphipoda (Crustacea) of Panama. Part III: parvorder Lysianassidira. ZooKeys 1216: 149-171. https://doi.org/10.3897/zookeys.1216.135258
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Abstract
Amphipods in the parvorder Lysianassidira are scavengers, often collected in sediment, coral rubble, algae, or among other invertebrates. Members of the parvorder have a head that is deeper than long, large coxae, lacinia mobilis present only on the left molar, and a mitten-shaped gnathopod 2 propodus with a long ischium. Nine species from two families within the parvorder are documented from Bocas del Toro, Panama. This research documents range extensions for eight species and an identification key to the species of Caribbean Lysianassidira of Panama is provided.
Resumen
Los anfípodos del parvorden Lysianassidira son carroñeros, a menudo recolectados en sedimentos, escombros de coral, algas o entre otros invertebrados. Los miembros del parvorden tienen una cabeza que es más profunda que larga, con coxas grandes, lacinia mobilis presenta solo en el molar izquierdo y un gnatópodo 2 en forma de manopla con un isquion largo. Nueve especies de dos familias dentro del parvorder están documentadas en Bocas del Toro, Panamá. Esta investigación documenta extensiones de rango para ocho especies y se proporciona una clave de identificación para las especies de Lysianassidira caribeña de Panamá.
Key words
Bocas del Toro, identification key, Lysianassidae, Lysianassoidea, Tryphosidae
Introduction
Parvorder Lysianassidira Dana, 1849 is comprised of 1243 species around the world, with several listed as incertae sedis (
Within the parvorder Lysianassidira, nine species of amphipods were collected from Bocas del Toro, Panama, with representatives from the families Lysianassidae and Tryphosidae. Regional diagnoses for each species collected during this study are provided herein. An identification key is provided to distinguish between the Lysianassidira species known from the Caribbean waters of Panama.
Methods
Coral rubble, sand, algae, and sponges were collected by hand and placed into buckets or plastic bags from various sites around Bocas del Toro, Panama at depths of 0–15 m. Coral rubble, sand, and algae were elutriated with freshwater to remove amphipods, and sponges were sorted through by hand. Live amphipods were sorted to morphospecies, placed in clove oil for imaging, and preserved in 99.5% EtOH for later examination. Preserved specimens were transferred to glycerol, measured from the tip of the rostrum to the base of the telson, and dissected under a stereomicroscope. Specimens were illustrated using a Meiji MT5900L phase contrast microscope with an Olympus U-DA drawing tube. Illustrations were digitally inked following
Results
Parvorder Lysianassidira Dana, 1849
Superfamily Lysianassoidea Dana, 1849
Family Lysianassidae Dana, 1849
Aruga
Diagnosis
Antenna 1 with strong callynophore in male and female. Antenna 2 flagellum elongate in male. Epistome not produced; upper lip produced. Maxilla 2 inner plate narrow. Gnathopod 1 simple. Gnathopod 2 minutely chelate. Uropod 2 inner ramus with dorsal notch, gradually narrowing distally. Uropod 3 outer ramus 2-articulate. Telson entire.
Aruga holmesi
Aruga holmesi
J.L. Barnard, 1955: 100, pls 27, 28; J.L.
Lysianopsis holmesi:
Lysianassa holmesi: J.L.
Material examined
Panama • 4.8 mm • 1 ♀; Bocas del Toro, Crawl Cay; 9.2376°N, 82.1438°W; depth 1.5–3 m, among coral rubble; 11 Aug 2021; K.N. White leg.;
Diagnosis
Upper lip projecting well beyond epistome; epistome concave. Gnathopod 1 propodus posterodistal margin slightly concave. Epimeron 3 posteroventral corner subquadrate, without tooth. Uropod 3 peduncle length at least 2 × width. Telson distal margin truncate, slightly emarginate, with two short setae on each side.
Distribution
USA: Folly Island, South Carolina; Florida from Perdido Key to the lower Florida Keys (
Ecology and remarks
These amphipods are associated with coral rubble and seagrass beds at depths of 1.5–120 m. Panamanian specimens agree closely with previous descriptions of the species.
Bonassa
Diagnosis
Antenna 1 with strong callynophore in male. Antenna 2 flagellum elongate in male. Epistome and upper lip produced. Maxilla 2 inner plate narrow. Gnathopod 1 simple. Gnathopod 2 minutely chelate. Uropod 2 inner ramus with dorsal notch, gradually narrowing distally. Uropod 3 outer ramus 1-articulate. Telson entire.
Bonassa bonairensis
Lysianassa
(?) bonairensis Stephensen, 1933a: 416–420, figs 1, 2;
Lysianassa bonairensis
J.L. Barnard, 1958: 94;
Bonassa bonairensis
Barnard & Karaman, 1991: 472;
Material examined
Panama • 2–3 mm • 1 ♀; Bocas del Toro, Swan Cay; 9.4533°N, 82.2983°W; depth 2–3 m, among algae; 4 Aug 2005; S. DeGrave leg.;
Diagnosis
Epistome produced, rounded, subequal to produced upper lip. Antenna 1 with strong callynophore in female. Gnathopod 1 propodus distally narrowing. Pereopod 7 basis greatly expanded, posteriorly rounded; merus greatly expanded, approximately 3 × width of carpus. Uropod 3 rami narrow, apically acute, and lacking plumose setae in female. Telson distal margin truncate, slightly emarginate.
Distribution
Lesser Antilles: Bonaire Island (
Ecology and remarks
These amphipods occur among algae and coral rubble at depths of 1–3 m. Panamanian specimens agree closely with previous descriptions of the species, with the exception of a slightly emarginate telson, with the exception of the uropod 3, which is documented for the first time in a female. This species is easily distinguishable based on the expanded pereopod 7 basis and merus. Panamanian specimens are a translucent white color when alive.
Concarnes
Diagnosis
Antenna 1 with strong callynophore in male, lacking in female. Antenna 2 flagellum short in male and female. Epistome and upper lip produced. Mouthparts forming quadrate bundle. Maxilla 2 inner plate broad. Gnathopod 1 simple. Uropod 2 inner ramus with dorsal notch, gradually narrowing distally. Uropod 3 outer ramus 2-articulate. Telson weakly cleft.
Concarnes concavus
Socarnes concavus
Shoemaker, 1933: 247–248, fig. 1; J.L.
Concarnes concavus
Barnard & Karaman, 1991: 477;
Material examined
Panama • 5–6 mm • 1 ♀; Bocas del Toro, Crawl Cay; 9.2475°N, 82.1290°W; depth 5 m, among coral rubble; 12 Aug 2021; K.N. White leg.;
Diagnosis
Head ocular lobe subacute. Epistome produced, rounded, subequal to produced upper lip. Gnathopod 1 basis slender, elongate; propodus distally narrowing. Gnathopod 2 minutely subchelate. Telson partially cleft, lobes apically rounded.
Distribution
USA: Santee River, South Carolina (
Ecology and remarks
These amphipods are associated with coral rubble and coarse sand at depths of 1–80 m. Panamanian specimens agree closely with previous descriptions of the species. This species is easily recognizable by the subacute ocular lobe, produced epistome and upper lip, and slender, elongate basis of gnathopod 1. Panamanian specimens have a distinct red coloration on the tips of antennae and on the anterior half of the body and have a white snowflake pattern on the posterior half of the body when alive.
Lysianopsis
Diagnosis
Antenna 1 with strong callynophore in male, weak or lacking in female. Antenna 2 flagellum short in male and female. Epistome not produced; upper lip produced. Maxilla 2 inner plate narrow. Gnathopod 1 simple. Gnathopod 2 minutely chelate. Uropod 2 inner ramus with dorsal notch, gradually narrowing distally. Uropod 3 outer ramus 1-articulate. Telson entire.
Lysianopsis hummelincki
Lysianassa hummelincki
Stephensen, 1933b: 438–440, fig. 1;
Lysianassa falcata
Stephensen, 1933b: 440–441, fig. 2;
Lysianopsis alba Barnard & Karaman, 1991: 499 (in part).
Falcanassa falcata Barnard & Karaman, 1991: 486.
Lysianopsis hummelincki Lowry & Stoddart, 1997: 82–89, figs 37–39.
Material examined
Panama • 4 mm • 1 ♂; Bocas del Toro, Hospital Point; 9.3320°N, 82. 2148°W; depth 1–3 m, among coral rubble; 22 June 2023; K.N. White leg.;
Diagnosis
Upper lip produced well beyond epistome; epistome straight. Gnathopod 1 of male prehensile. Pereopod 7 basis slightly expanded, posterior margin almost straight, merus slightly expanded, approximately 1.4 × width of carpus. Uropod 3 peduncle length about 1.5 × width; outer ramus 1-articulate. Telson distal margin rounded.
Distribution
Lesser Antilles: Curaçao (
Ecology and remarks
These amphipods are associated with sand and coral rubble at depths of intertidal 0–12 m. Panamanian specimens agree closely with previous descriptions with the exception of the almost straight posterior margin on the pereopod 7 basis, which was described by
Lysianopsis ozona
Lysianopsis ozona Lowry & Stoddart, 1997: 87–91, figs 40–42.
Material examined
Panama • 3.2–8.5 mm • 2 ♀; Bocas del Toro, Bastamientos; depth 0–1 m, mangrove scrapings; 1 Aug 2005; T.A. Haney leg.;
Diagnosis
Epistome concave, subequal to upper lip. Gnathopod 1 propodus posterodistal margin straight; not sexually dimorphic. Uropod 3 peduncle length approximately 1.5 × width; outer ramus 2-articulate. Telson apical margin slightly truncate, apical margin with four short setae medially.
Distribution
USA: Eastern Gulf of Mexico (
Ecology and remarks
These amphipods are associated with sand, coral rubble, and various invertebrates at depths of 0–29 m. Panamanian specimens agree closely with the description provided by
Shoemakerella
Diagnosis
Antenna 1 with weak callynophore in male, lacking in female. Antenna 2 flagellum short in male and female. Epistome not produced; upper lip produced. Maxilla 2 inner plate wider than outer plate. Gnathopod 1 simple. Pereopods 3–4 merus not enlarged compared to carpus. Uropod 2 inner ramus with dorsal notch, abruptly narrowing distally. Uropod 3 outer ramus 1-articulate. Telson entire, dorsal setae inserted proximally (compared to other genera).
Shoemakerella cubensis
Lysianax cubensis
Stebbing, 1897: 29–30, pl. 7B;
Lysianassa cubensis
Stebbing, 1906: 38;
Lysanopsis alba
Pearse, 1912: 369, fig. 1 (in part);
Shoemakerella nasuta
Pirlot, 1936: 265–266;
Lysianopsis cubensis Hurley, 1963: fig. 21a.
Lysianassa nasuta
Ortiz, 1978: 8;
Shoemakerella cubensis
Barnard & Karaman, 1991: 530;
Material examined
Panama • 1.5–4 mm • 3 ♀, 1 juvenile; Bocas del Toro, Hospital Point; 9.3336°N, 82.2188°W; depth 15 m, among coral rubble and Halimeda; 6 Aug 2005; S. DeGrave and M. Salazar leg.;
Diagnosis
Head and body with tiny setules. Epistome strongly concave. Pereopod 6 basis posterior margin nearly straight. Pereopod 7 propodus length ~5 × width. Telson apex rounded.
Distribution
USA: Panama City to Dry Tortugas, Florida (
Ecology and remarks
These amphipods are associated with algae and coral rubble at depths of 2–69 m. Panamanian specimens closely resemble previously described specimens and can be readily distinguished from Shoemakerella lowryi Gable & Lazo-Wasem, 1990 based on the pereopod 6 basis posterior margin, pereopod 7 propodus length relative to the carpus length, and the telson apex. Panamanian specimens are yellow-orange in color when alive.
Shoemakerella lowryi
Lysianassa punctata
Kunkel, 1910: 8–10, fig. 1;
Shoemakerella lowryi Gable & Lazo-Wasem, 1990: 727–733, figs 5–7.
Material examined
Panama • 2–5.5 mm • 1 ♂; Bocas del Toro, San Cristobal; 9.2625°N, 82.2350°W; depth 15 m, among coral rubble; 10 August 2021; K.N. White leg.;
Diagnosis
Head and body with tiny setules. Epistome weakly concave. Pereopod 6 basis posterior margin slightly concave. Pereopod 7 propodus length ~9 × width. Telson apex truncate.
Distribution
Bermuda (
Ecology and remarks
These amphipods are associated with algae, seagrass, and coral rubble at depths of 0.5–9 m. Panamanian specimens closely resemble previously described specimens and can be readily distinguished from Shoemakerella cubensis based on the pereopod 6 basis posterior margin, pereopod 7 propodus length relative to the carpus length, and the telson apex. Panamanian specimens are transparent white in color with brown spots when alive.
Family Tryphosidae Lowry & Stoddart, 1997
Lepidepecreum
Diagnosis
Antenna 1 with strong callynophore in male and weak callynophore in female. Antenna 2 of male elongate. Antenna 2 peduncular article 3 elongate in male and female. Maxilla 2 inner plate narrow. Gnathopod 1 subchelate; coxa large, about as long as coxa 2; carpus long (length 2 to 4 × width). Uropod 2 inner ramus without distinct dorsal notch. Uropod 3 outer ramus 2-articulate. Telson cleft.
Lepidepecreum cf. magdalenensis
Orchomenella magdalenensis Shoemaker, 1942: 4–7, fig. 1.
Lepidepecreum magdalenensis
Lowry & Stoddart, 2002: 173–174;
Material examined
Panama • 2–3 mm • 6 ♂, 16 ♀; Bocas del Toro, Drago Beach; 9.4172°N, 82.3248°W; depth 0–1 m, in sand; 27 June 2023; K.N. White leg.;
Diagnosis
Head ocular lobe subrectangular. Gnathopod 1 carpus as long as propodus. Epimeron 3 posteroventral corner subquadrate. Urosomite 1 with dorsodistally acute carina. Uropod 3 inner ramus with two marginal spines.
Distribution
USA: Pacific California (
Ecology and remarks
These amphipods are associated with sand at depths of 0.5–27 m. Panamanian specimens closely resemble previously described specimens, except for a weak callynophore in females (strong in original description) and uropod 3 inner ramus having 2 marginal spines (3 in original description).
Orchomenella
Diagnosis
Antenna 2 of male flagellum elongate. Antenna 2 peduncular article 3 short. Maxilla 2 inner plate narrow. Gnathopod 1 subchelate; carpus short (length less than 2 × width). Uropod 2 inner ramus without distinct dorsal notch. Telson cleft.
Orchomenella thomasi
Orchomenella thomasi
Lowry & Stoddart, 1997: 109–113, figs 52–53;
Material examined
Panama • 1.5 mm • 1 ♀; Bocas del Toro, Cayo Zapatilla 1; 9.2700°N, 82.0587°W; depth 10–11 m, among coral rubble; 28 June 2023; K.N. White leg.;
Diagnosis
Head ocular lobe subtriangular. Gnathopod 1 carpus shorter than propodus. Epimeron 3 posteroventral corner acute. Urosomite 1 with dorsodistally acute carina. Uropod 3 inner ramus bare; outer ramus 2-articulate.
Distribution
Ecology and remarks
These amphipods are associated with sand and coral rubble at depths of 10–73 m. Panamanian specimens closely resemble previously described specimens. Panamanian specimens are white in color when alive.
Identification Key to the Caribbean Lysianassidira of Panama
| 1 | Eye absent; pereopod 5 basis narrowly expanded | 2 |
| – | Eye present, well developed; pereopod 5 basis broadly expanded | 3 |
| 2 | Head ocular lobe produced; epimeron 3 posteroventral margin with acute tooth; telson deeply cleft, about 75% | Paracentromedon carabicus |
| – | Head ocular lobe evenly rounded; epimeron 3 posteroventral margin rounded; telson shallowly cleft, less than 50% | Vemana compressa |
| 3 | Gnathopod 1 subchelate; urosomite 1 with dorsodistally acute carina; uropod 2 inner ramus without distinct dorsal notch (Figs |
4 |
| – | Gnathopod 1 simple; urosomite 1 without dorsodistal carina; uropod 2 inner ramus with distinct dorsal notch (Fig. |
5 |
| 4 | Antenna 2 peduncle article 3 long in female; head ocular lobe subrectangular; gnathopod 1 carpus as long as propodus; epimeron 3 posteroventral corner subquadrate; uropod 3 inner ramus with marginal spines (Fig. |
Lepidepecreum magdalenensis |
| – | Antenna 2 peduncle article 3 short in female; head ocular lobe subtriangular; gnathopod 1 carpus shorter than propodus; epimeron 3 posteroventral corner acute; uropod 3 inner ramus bare (Fig. |
Orchomenella thomasi |
| 5 | Gnathopod 1 dactylus reduced, complex, covered in long, slender cuticular teeth; telson entire | Eclecticus eclecticus |
| – | Gnathopod 1 dactylus not reduced, simple; telson entire or partially cleft | 6 |
| 6 | Gnathopod 2 minutely subchelate; telson partially cleft (Fig. |
Concarnes concavus |
| – | Gnathopod 2 minutely chelate (Figs |
7 |
| 7 | Maxilla 2 inner plate wider than outer plate (Fig. |
8 |
| – | Maxilla 2 inner plate narrow, similar in width to outer plate (Fig. |
9 |
| 8 | Pereopod 6 basis posterior margin nearly straight; pereopod 7 propodus length ~5 × width; telson apex rounded (Fig. |
Shoemakerella cubensis |
| – | Pereopod 6 basis posterior margin slightly concave; pereopod 7 propodus length ~9 × width; telson apex truncate (Fig. |
Shoemakerella lowryi |
| 9 | Epistome rounded; uropod 3 outer ramus 1-articulate (Fig. |
10 |
| – | Epistome concave; uropod 3 outer ramus 2-articulate (Fig. |
11 |
| 10 | Epistome produced, subequal to produced upper lip; gnathopod 1 basis slender; pereopod 7 basis greatly expanded, posteriorly rounded, merus greatly expanded, approximately 3 × width of carpus (Fig. |
Bonassa bonairensis |
| – | Epistome not produced, upper lip produced; gnathopod 1 basis stout; pereopod 7 basis slightly expanded, posterior margin almost straight, merus slightly expanded, approximately 1.4 × width of carpus (Fig. |
Lysianopsis hummelincki |
| 11 | Upper lip projecting well beyond epistome; gnathopod 1 propodus posterodistal margin slightly concave; uropod 3 peduncle long, length at least 2 × width; telson apical margin slightly emarginate (Fig. |
Aruga holmesi |
| – | Upper lip subequal to epistome; gnathopod 1 propodus posterodistal margin straight; uropod 3 peduncle short, length approximately 1.5 × width; telson apical margin slightly truncate (Fig. |
Lysianopsis ozona |
Discussion
The results of this study represent range extensions for eight species of lysianassid amphipods to include the Caribbean waters of Panama. One species collected in this study, Concarnes concavus, has been recorded from the Caribbean of Panama by
Characters that have been used to identify lysianassid amphipods in the past, such as setae patterns on the dorsal surface of the body appear to be variable in Panamanian specimens and should not be used for identification. Sexual dimorphism is also used frequently but can be problematic when you have only one specimen or gender. Mouthparts are also often used as diagnostic characters which can be difficult for non-experts; thus, I included as many other characters as possible in this identification key.
Acknowledgements
Logistical support and facilities were provided by Georgia College & State University Department of Biological and Environmental Sciences and the Smithsonian Tropical Research Institute (STRI). Special thanks to Carolina Cesar and Valentina Cardona for assistance with diving and collecting in Bocas del Toro. The author also wishes to thank Dr Lauren Hughes and the Amphipod Taxonomy Course members for collecting assistance in 2023. Special thanks go to Sara LeCroy for her loan of specimens collected in 2005 and to Sally Sir, Sara LeCroy, and Tammy Horton for suggestions to improve the manuscript.
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Funding
Funding for this study was provided by a National Science Foundation grant: Collaborative Research: ARTS: Understanding Tropical Invertebrate Diversity Through Integrative Revisionary Systematics and Training (1856421).
Author contributions
Conceptualization: KNW. Data curation: KNW. Formal analysis: KNW. Funding acquisition: KNW. Investigation: KNW. Methodology: KNW. Project administration: KNW. Writing - original draft: KNW. Writing - review and editing: KNW.
Author ORCIDs
Kristine N. White https://orcid.org/0000-0002-5203-1656
Data availability
All of the data that support the findings of this study are available in the main text or Supplementary Information.
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Supplementary material
Locality table
Data type: xlsx