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418
Accepted by J. Goy: 26 Jul. 2018; published: 25 Oct. 2018
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4504 (3): 418
–
430
http://www.mapress.com/j/zt/
Article
https://doi.org/10.11646/zootaxa.4504.3.7
http://zoobank.org/urn:lsid:zoobank.org:pub:9FC7CB81-1174-469F-8E70-01447A1D9BA1
A new species of Alvinocaris (Crustacea: Decapoda: Caridea: Alvinocarididae)
from Costa Rican methane seeps
JOEL W. MARTIN
1,4
, ADAM R. WALL
1
, TIM SHANK
2
, HARIM CHA
3
,
CHARLOTTE A. SEID
3
& GREG W. ROUSE
3,4
1
Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, CA 90007
2
Biology Department MS #33, Woods Hole Oceanographic Institution, Woods Hole, MA 02543
3
Scripps Institution of Oceanography, UCSD, La Jolla, CA 92093-0202
4
Corresponding author. E-mail: jmartin@nhm.org, grouse@ucsd.edu
Abstract
A new caridean shrimp, Alvinocaris costaricensis, is described from methane seeps in the eastern Pacific off Costa Rica.
The new species is the 16
th
described species of the genus, and by molecular analysis appears closest to Alvinocaris komaii
from the Lau Basin, southwestern Pacific, but shares certain morphological characters with A. lusca from the Galapagos
Rift and A. muricola from the West Florida Escarpment, as well as with A. kexueae from the Manus Basin in the Southwest
Pacific.
Key words: Caridea, Alvinocaris, methane seeps, Costa Rica
Introduction
The caridean shrimp family Alvinocarididae is an unusual and interesting assemblage of species known only from
hydrothermal vents and cold seeps (Martin & Haney 2005; De Grave & Fransen 2011). Currently, the family
consists of nine genera and at least 32 species (Komai & Segonzac 2005; De Grave & Fransen 2011; Yahagi et al.
2014; Komai & Tsuchida 2015; Komai et al. 2016) (Table 1). More undescribed species are known to exist in
various collections around the world (e.g. see Komai & Segonzac 2005), and it is common to encounter
undescribed species as new vent and seep sites are explored. The most geographically widespread and speciose
genus, Alvinocaris, was reviewed by Komai & Segonzac (2005), at which time there were eight described species.
Additional species have been described in the twelve years since their review (listed in De Grave & Fransen 2011;
see also Yahagi et al. 2014; Vereschaka et al. 2015; Wang & Sha 2017; Table 2). Below, we describe a new species
of Alvinocaris from relatively shallow methane seeps off the coast of Costa Rica in the eastern Pacific Ocean.
TABLE 1. Shrimp genera of the family Alvinocarididae as of mid-2018.
Genus No. of described species
Alvinocaridinides Komai & Chan, 2010 2
Alvinocaris Williams & Chace, 1982 16 (including this paper)
Chorocaris Martin & Hessler, 1990* 5
Manuscaris Komai & Tshuchida, 2015 1
Mirocaris Vereschaka, 1997 2
Nautilocaris Komai & Segonzac, 2004 1
Opaepele Williams & Dobbs, 1995 3
Rimicaris Williams & Rona, 1986 2
Shinkaicaris Komai & Segonzac, 2005 1
*suggested to be synonymous with Rimicaris by Vereschaka et al. (2015)
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A NEW SPECIES OF ALVINOCARIS FROM COSTA RICAN
TABLE 2 . Species of the genus Alvinocaris described as of mid-2018, in alphabetical order, and the general geographic
region in which they were discovered.
A. alexander Ahyong, 2009 southern Kermadec Ridge, southwest Pacific
A. brevitelsonis Kikuchi & Hashimoto, 2000 Minami-Ensei Knoll, Okinawa Trough
A. chelys Komai & Chan, 2010 Gueishandao, Taiwan
A. costaricensis (this paper) Costa Rica, eastern Pacific
A. dissimilis Komai & Segonzac, 2005 Minami-Ensei Knoll, Okinawa Trough
A. komaii Zelnio & Hourdez, 2009 Lau Basin, southwest Pacific
A. kexueae Wang & Sha, 2017 Manus Basin, southwest Pacific
A. longirostris Kikuchi & Ohta, 1995 Okinawa Trough
A. lusca Williams & Chace, 1982 Galapagos Rift, Rose Garden
A. markensis Williams, 1988 Mid-Atlantic Ridge
A. methanophila Komai, Shank & Van Dover, 2005 Blake Ridge
A. muricola Williams, 1988 West Florida Escarpment
A. niwa Webber, 2004 Rumble V seamount, Pacific
A. solitaire Yah a g i et al., 2014 Central Indian Ridge, Indian Ocean
A. stactophila Williams, 1988 North Central Gulf of Mexico
A. williamsi Shank & Martin, 2003 Menez Gwen, North Atlantic
Materials and methods
Specimens were collected from off the Pacific coast of Costa Rica during two cruises (AT 15-44 and 15-59 in 2009
and 2010, respectively) to study methane seeps, with sampling carried out via the Deep Submergence Vehicle
(DSV) Alvin. Shrimp were collected via the DSV Alvin and photographed alive onboard ship with a Canon
PowerShot G9 camera mounted on a Leica S8APO stereomicroscope.
Specimens for morphological study were fixed in 10% seawater-formaldehyde and then preserved in 70%
ethanol after rinsing. Two specimens from the 2009 expedition, both from Mound 12 (details below), were sent to
the first author for initial examination and illustration. All illustrations in this paper are of the larger (female) of
those two specimens, which is designated the holotype. All illustrations were made with the aid of a Wild M5APO
dissecting microscope and drawing tube.
Abbreviations used in the description are CL (carapace length) and TL (total length), and OCL (orbital
carapace length), and for institutions NHMLAC (Natural History Museum of Los Angeles County), MZUCR
(Museo de Zoología, Universidad de Costa Rica), and SIO-BIC (Benthic Invertebrate Collection of the Scripps
Institution of Oceanography)
Specimens for DNA analysis were preserved directly in 95% ethanol, and genomic DNA was extracted from
legs of seven specimens of the new species using a Qiagen DNeasy Tissue kit. Cytochrome oxidase subunit 1
(COI) (partial, approximately 700 bp) was amplified using the primer pair LCO1490 (5’-GGT CAA CAA ATC
ATA AAG ATA TTG G- 3’) and HCO2198 (5’-ACT TCA GGG TGA CCA AAA AAT CA-3’) (Folmer et al.
1994). PCR mixtures contained 12.5 μL ProMega GoTaq Green DNA polymerase (3mM MgCl2, 400 μM each
dNTP, 1U Taq), between 50–100 ng DNA, and a reaction profile of 94ºC for 180s, 5 cycles of 94°C for 30s, 47°C
for 45s, and 72°C for 60s, 30 cycles of 94°C for 30s, 52°C for 45s, and 72°C for 60s and finally 72°C for 300s was
used. The COI sequences, lacking indels, were unambiguously aligned and blasted to other alvinocarids. TCS
(Clement et al. 2000) was used to construct a COI haplotype network for the seven specimens of A. costaricensis n.
sp.
The seven new sequences (GenBank Accession Numbers MH645099-MH645105) were also combined with a
COI dataset of available species of Alvinocarididae in GenBank, largely following those used in Vereshchaka et al.
(2015), including their choice of outgroup, the acanthephyrid shrimp Acanthephyra purpurea. COI sequences were
managed in Mesquite (Maddison & Maddison 2011) and aligned using Muscle (Edgar 2004). Pairwise
comparisons of COI sequences were conducted, and uncorrected distances were estimated in PAUP* 4.0b15
(Swofford 2002). The sequences were analyzed using maximum parsimony (MP) in PAUP* and maximum
likelihood (ML) with RaxML GUI and RaXML 8 (Silvestro & Michalak 2012; Stamatakis 2014). MP analyses
were run using heuristic searches (100 random stepwise addition replicates; bisection re-connection (TBR); zero-
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length branches collapsed). Clade support was assessed using jackknifing (37% deletion) with 1000
pseudoreplicates with 100 random additions per iteration. The ML analysis was run with the data partitioned by
codon using the GTR+Γ model of substitution. Bootstrap support values were estimated by 1000 standard
bootstrap pseudo-replicates with the same model and partitions.
Taxo n o m y
Alvinocarididae
Alvinocaris costaricensis, new species
Material examined. Holotype: SIO-BIC C12202, female (TL ~42 mm, CL 17.4 mm, OCL 10.4 mm, carapace
height 6.0 mm), Eastern Pacific Ocean, Costa Rica, Mound 12, Alvin dive 4503, 8.9307° N, 84.3072° W, 1005 m,
collector Greg Rouse, Feb/24/2009. Paratypes: SIO-BIC C13298, SIO-BIC C13299, same collecting data as for
holotype; MZUCR-3569-01, same collecting data as for holotype. NHMLAC LACM CR 2009.1 (ex SIO-BIC
C11140-1), ovigerous female (OCL 17.6 mm, TL 55 mm), same collecting data as for holotype. SIO-BIC C11157
and C12203, Eastern Pacific Ocean, Costa Rica, Mound 12, Alvin dive 4511, 8.9305° N, 84.3123° W, 1001 m,
collector Greg Rouse, March/5/2009; SIO-BIC C12203 1 badly damaged specimen (TL 13.2 mm, OCL 4.9 mm;
abdomen disarticulated, sex undetermined). Although damaged, this specimen was sequenced and is the same
species. SIO-BIC C11157 was sequenced and is the same species. SIO-BIC C11209, sex not determined, (TL 77.6
mm, CL 30.4 mm, OCL 17.1 mm; left side of carapace slightly inflated), Eastern Pacific Ocean, Costa Rica, Jaco
Scarp, Alvin dive 4590, 9.1176° N, 84.8395° W, 1800 m, collector Greg Rouse, Jan/11/2010. SIO-BIC C11183,
C11186, sex not determined, Eastern Pacific Ocean, Costa Rica, Mound 12, Alvin dives 4587 and 4588, 8.9307° N,
84.3072° W, 1005 m collector Greg Rouse, Jan/08 and 09/2010. These specimens were sequenced and are the same
species.
Description. Body relatively robust for the genus, integument thin, smooth, shiny. Carapace (Figs. 1, 2) with
strong suborbital spine (Fig. 3A) and well developed pterygostomial spine exceeding length of orbital spine,
otherwise unarmed.
Rostrum (Figs. 1B, 2, 3A) well developed (tip broken in holotype but well developed in several paratypes),
extending forward and only slightly downward, with strong, sharp, anterior-curving teeth in row on dorsal border,
extending backward to about, or slightly posterior to, midlength of carapace, bearing 5–7 weakly developed teeth
along ventral border. Weak dorsal carina extending backward from rostrum along carapace to posterior border.
Larger specimens with rostrum strongly up-turned distally, bearing up to 10 dorsal and forward-directed teeth,
exceeding anteriorly beyond tip of scaphocerite and antennular peduncle.
Eighth thoracic sternite with well-developed and acute median tooth directed anteroventrally, produced beyond
coxa of pereopod 5 and visible in lateral view (Fig. 2).
Abdomen (Figs. 1A, 2) well developed, somite 6 with sharp posteriorly directed tooth on either side extending
posteriorly along telson (Fig. 3D); somite 5 with similar but shorter tooth and with acute posteroventral border;
somite 4 with acute posteroventral border.
Telson (Fig 2, 3D, E) long, exceeding length of uropods, lateral margins straight, slightly converging
posteriorly; each lateral margin bearing row of 7 movable spines; posterolateral corners each with single large
slightly medially-curved spine; posterior border bearing two pairs of small spines and 4 plumose setae; posterior
border slightly indented as shown. Uropodal rami each with border of plumose setae; exopod broader than
endopod, bearing diaresis and single lateral spine just posterior to acute tooth at border of diaresis, as shown (Fig.
3D).
Eyes fused mesially but distinct, each with small anterodorsal tubercle (Fig. 3A). Cornea with diffuse
pigmentation internally but no clear pigmented layer or region.
Antennular peduncle (Fig. 3C) extending beyond antennal scale, with relative lengths of articles 1 > 2 > 3;
article 1 with large lateral spine extending clearly beyond similar spine of article 2. Antennal scale (Fig. 3B) broad,
distally rounded, with acute anterolateral tooth extending almost to full length of scale.
Mouthparts (Fig. 4A–F) typical of genus (Komai & Segonzac 2005; Vereschaka et al. 2015).
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A NEW SPECIES OF ALVINOCARIS FROM COSTA RICAN
FIGURE 1. A, Alvinocaris costaricensis new species, holotype female, SIO-BIC C12202, Eastern Pacific Ocean, Costa Rica,
live color photograph. B, same species, paratype female, C11186.
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FIGURE 2. Alvinocaris costaricensis new species, holotype female, SIO-BIC C12202, Eastern Pacific Ocean, Costa Rica,
lateral view, extremity of rostrum broken. Carapace length 10.4 mm (measured from back of the orbital sinus in a straight line
to posterior border of carapace).
First pereopod (cheliped) (Figs. 1A, 2, 3F–H) well developed, long, extending beyond bases of antennae when
outstretched; chela delicate for genus and strongly curved inward, bearing delicate pectinations along cutting
borders of both fingers; fingers of chela approximately 1.5 times length of propodal palm; carpus cup-shaped to
receive proximal end of propodus, and bearing brush of “cleaning setae” and large, acute tooth on inner margin.
Second pereopod (Figs. 2, 3I–K) shorter than first; chela delicate, approximately 4 times longer than wide,
both fingers bearing pectinate setal borders on cutting margin; fingers slightly exceeding length of palm; ischium
with single movable ventral spine.
Third through fifth pereopods (Fig. 5) similar, long and delicate; propodus longer than merus; merus longer
than carpus. Dactylus short, stout, with recurved sclerotized tip and ventral single row of 4 or 5 short, sclerotized
spines. Propodus with row of regularly spaced short spines along ventral border. Merus with 2 large, ventral
movable spines on P3, none on P4 and P5. Ischium with 2 ventral movable spines on P3 and P4, none on P5.
Female pleopod (Fig. 4G) with subequal rami bearing plumose setae; appendix interna short, simple, tapering
distally.
Coloration. Photographs of the holotype (Fig. 1A) and one paratype (Fig. 1B) show a largely translucent to
white shrimp with a pale or beige carapace, some orange or reddish coloration on the mouthparts, and delicate red
reticulations on the carapace and abdomen. The eye, although reflecting light in the photographs, appears to be
orange-red.
Etymology. The specific epithet reflects the location of the methane seeps off the Pacific coast of Costa Rica.
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FIGURE 3. Alvinocaris costaricensis new species, holotype female, SIO-BIC C12202, Eastern Pacific Ocean, Costa Rica. A,
eye and section of rostrum (above) and orbital spine (below); note minute dorsal tubercle on eye. B, antennal scale, right side,
dorsal view. C, antennular peduncle, right side, dorsal view. D, telson and left uropods, dorsal view. E, extremity
(approximately distal half) of telson. F, pereopod 1 (cheliped), right side, ventro-lateral view. G, pereopod 1, dorsal view. H,
higher magnification of pereopod 1, dorsal inner view. I, second pereopod, right side, lateral view. J, pereopod 2 propodus and
chela, outer view. K, pereopod 2 chela, inner view.
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FIGURE 4. Alvinocaris costaricensis new species, holotype female, SIO-BIC C12202, Eastern Pacific Ocean, Costa Rica. A,
left mandible, inner view. B, maxilla, inner view. C, maxilliped 1. D, maxilliped 2. E, maxilliped 3, lateral view. F, maxilliped 3
dactylus, inner view. G, first pleopod and appendix interna, right side.
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FIGURE 5. Alvinocaris costaricensis new species, holotype female, SIO-BIC C12202, Eastern Pacific Ocean, Costa Rica. A,
pereopod 3, left side, lateral view. B, same, close up of dactylus and distal half of propodus. C, pereopod 4, left side, lateral
view. D, close up of dactylus and distal half of propodus. E, pereopod 5, left side, lateral view. E, close up of dactylus and distal
half of propodus.
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FIGURE 6. Relationships among selected members of Alvinocarididae based on COI sequence data. A, Maximum likelihood
topology; numbers at nodes indicate bootstrap values. B, one of eight shortest trees from the maximum parsimony analysis of
COI matching jackknife consensus tree; numbers at nodes refer to parsimony jackknife score. C. Haplotype network generated
by TCS (Clement et al. 2002); holotype and 5 paratypes have identical COI sequences; paratype C11186 differs by one base.
Generic type species indicated (Type). Seep-dwelling species indicated (S).
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DNA sequence analysis. The seven COI sequences for A. costaricensis n. sp. were very similar, with six
(including that of the holotype) being identical, and one haplotype differing by only one base (Fig. 6C). These had
a closest pairwise distance (of ~12%) to Alvinocaris komaii (GenBank KP759373) from West Pacific hydrothermal
vents. Of the 505 bases in the COI dataset, 323 were constant, 158 were parsimony-informative, and 24 were
variable but parsimony-uninformative. The ML analysis (Fig. 6A) recovered Alvinocaris as paraphyletic, with A.
costaricensis n. sp. as the sister group to A. komaii. The Rimicaris+Opaepele+Shinkaicaris clade was nested within
Alvinocaris as sister to the A. costaricensis n. sp. + A. komaii clade, though this, and most other deeper nodes, had
low support. The four-known seep-dwelling alvinocaridids, A. methanophila, A. muricola, A. stactophila and A.
costaricensis n. sp., were scattered across the phylogeny and all had hydrothermal vent-dwelling taxa as sister
groups. The MP analysis (Fig. 6B) found eight shortest trees of length 547. These eight trees varied only slightly;
one of the most parsimonious trees with the same topology as the majority-rule consensus tree, and that of the of
majority-rule consensus tree for the parsimony jackknife analysis, is shown in Fig. 6B. The MP analysis recovered
a monophyletic Alvinocaris (with the exception of A. methanophila), though with poor support; as in the ML
results, A. costaricensis n. sp. was sister group to A. komaii.
Remarks. Morphological comparison. Alvinocaris costaricensis n. sp. is a large species for an alvinocaridid,
with some paratypes (e.g. SIO-BIC C11209 from the Jaco Scarp site) exceeding 75 mm TL and 64 mm OCL. The
species is easily identifiable as a member of Alvinocaris based on several characters uniquely shared by members
of that genus: the laterally compressed, well-developed and toothed rostrum, dentate posterior border on abdominal
somite 4, minute dorsal tubercle on the surface of the eye, strongly curved and minutely pectinate chelae, small
ischial spine on the second pereopod, parallel rows of small spines on the telson, and spination of pereopods 3 and
4, among other characters (Komai & Segonzac 2005, Vereschaka et al. 2015).
Among species of Alvinocaris, the new species is similar to A. muricola and also to A. kexueae in the size
(length) of the rostrum, the orientation of the dorsal rostral spination, and the relatively shallow dorsal angle of the
carapace. In all three species, the rostrum slopes more gradually, and bears larger teeth, than in most Alvinocaris
species (Komai & Segonzac 2005). However, there is much morphological variation in the rostrum of A. muricola.
Using the diagnostic key to species provided by Komai & Segonzac (2005), the new species keys to Alvinocaris
lusca, another eastern Pacific (Galapagos Rift and East Pacific Rise) species, based on the number of ventral rostral
teeth (fewer than 5) and the length/width ratio of the antennal scale. Additionally, A. lusca is one of the few species
of Alvinocaris that shares with the new species, a slightly indented posterior border of the telson (though not as
indented as in A. costaricensis). However, A. costaricensis differs from A. lusca in having a far slenderer major
cheliped, 2 (rather than 3) ventral meral spines on pereopod 3, and a much longer lateral spine on the basal article
of the antennal peduncle, which barely exceeds the length of the spine of the second article in A. lusca.
An interesting departure from typical species of Alvinocaris is the absence of ventral meral spines on pereopod
4. Komai and Segonzac (2005) considered the presence of these spines on pereopods 3 and 4 diagnostic for the
genus Alvinocaris. The new species has prominent meral spines on P3, but not on P4 or P5 indicating that the
generic diagnosis requires slight amendment. This difference does not warrant the erection of another genus, as all
other characters fit well within the known range of features described for the other 16 species of Alvinocaris. The
indented terminal border of the telson is also unique among species in the genus, with A. lusca being the closest
match in that character. However, variation in this character is known (Komai & Segonzac 2005), so we hesitate to
suggest this as a determining feature for field identification.
Molecular comparison. We place the new species in Alvinocaris, even though there is evidence from the ML
analysis (albeit weakly supported) (Fig. 6A) that the genus, as currently construed, could be paraphyletic (and
polyphyletic when A. methanophila is considered). However, further sequence data are required to properly resolve
this situation, and that question is beyond the scope of this study. Vereshchaka et al. (2015) also found a
paraphyletic Alvinocaris based on their analysis of COI and also of the available 16S rDNA sequences, but
maintained the genus as currently formulated, although they suggested that the sequence for A. methanophila on
GenBank (AY163260) was the result of incorrect identification or processing of the material. Vereshchaka et al.
(2015) did recover Alvinocaris (including A. methanophila) as a well-supported clade based on morphology, and A.
costaricensis n. sp. shared these features (notably the laterally compressed and carinate rostrum, eyes fused
medially with each bearing a small tubercle, and paired dorsal spines on the telson).
Co-occurring species. Another, and smaller, alvinocaridid species, differing both morphologically (with a
shorter rostrum lacking the extensive dorsal teeth seen in A. costaricensis) and molecularly, was also collected
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from the Mound 12 site (AD 4501, 1008 m depth). Further specimens of that species in the SIO collection represent
yet another undescribed species.
TABLE 3 . Localities and GenBank Accession numbers for alvinocarid shrimp used in this study.
*Formerly in Chorocaris, synonymized with Rimicaris by Vereshchaka et al. (2015).
Acknowledgements
The authors are grateful to Chief Scientist Lisa Levin, the captain and crew of the RV Atlantis, the crew of the DSV
Alvin, and the scientific participants of AT 15-44 and 15-59 for crucial assistance in specimen collection.
Collection and analysis of the Costa Rica specimens for this project was funded by US National Science
Foundation grants to L. Levin and GWR (OCE-0826254 and OCE-0939557). JWM and AW thank the Natural
History Museum of Los Angeles County for support.
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