Synalpheus
Synalpheus is a genus of snapping shrimp of the family Alpheidae, presently containing more than 100 species; new ones are described on a regular basis, and the exact number even of described species is disputed.
''Zuzalpheus''
The genus Zuzalpheus was established for S. gambarelloides, S. brooksi, and their closest relatives, which contain several notably eusocial species. While these do seem to form a clade, it is not fully resolved whether or not they are indeed the sister taxon of all the remaining Synalpheus.However, a detailed cladistic study of morphological characters found well marked differences between the proposed two genera and concluded that the supposed species groups around S. biunguiculatus/S. coutierei, S. brevicarpus and S. neomeris are neither clearly defined nor, as it seems, monophyletic, while the group separated in Zuzalpheus was clearly distinct in characters of the minor first walking legs, and usually distinct in some others; in all Synalpheus sensu stricto checked to date, the transverse setal comb on the back of the minor first pereopod dactyl is missing, and the carpus is plump and small.
Moreover, their stylocerite clearly extends beyond the whole first segment of the first antennae, their scaphocerite blade is never missing, the fixed finger of the major first pereopod is about the same length as the dactyl and the uropodal exopod always has one tooth only; these traits however may be also present in some of the species separated in Zuzalpheus though as far as can be told, most of these differ. While the authors of the 2008 analysis made no explicit comment on the status of the newly proposed genus, they thus found the S. gambarelloides species group to be well-marked, and their results certainly harden the case for recognition of Zuzalpheus at least as a subgenus. However, this analysis only included 2 species from the S. gambarelloides species group, and none from the Caribbean, where the vast majority of the group resides. Opinion is still divided, however, and recently Zuzalpheus was synonymised with Synalpheus.
Distribution
In the narrower sense, Synalpheus occur in the East Pacific where they are most plentiful and probably originated, and to a lesser extent in the Atlantic and the Indian Ocean; the species placed in Zuzalpheus occur mainly in the western Atlantic where their lineage probably originated, and to a lesser extent in the eastern Atlantic and Indian Ocean, and the East Pacific. It may thus be that the closure of the Isthmus of Panama in the Piacenzian was a key factor in separating the two lineages, as species referred to Synalpheus sensu stricto are most plentiful in the western Pacific.Snapping
The snapping behaviour of Synalpheus is rather well studied. In Synalpheus parneomeris, peak to peak source levels of 185–190 dB re 1 μPa at 1 m were measured, depending on the size of the claw.Eusociality
The only known eusocial aquatic species occur within the genus Synalpheus. The species known to be eusocial are S. brooksi, S. chacei, S. elizabethae, S. filidigitus, S. rathbunae, S. regalis, S. microneptunus, and S. duffyi as well as potentially S. riosi. Eusociality has evolved at least three times within Synalpheus. It appears that there were multiple rapid radiations between 3 and 9 mya from which the ancestors of these eusocial species appeared. Eusociality is thought to have arisen due to competition for space, because among the species that host Synalpheus, empty sponges are rarely found. It also appears that kin selection was necessary for this evolution to occur because the only species in which eusociality has appeared are non-dispersing shrimp that hatch directly into crawling individuals. Until recently, eusocial species of Synalpheus have appeared in far greater abundance than and appear to outcompete less social species for space in sponges.Species
Synalpheus contains the following species:- Synalpheus agelas Pequegnat & Heard, 1979
- Synalpheus albatrossi Coutière, 1909
- Synalpheus amabilis De Man, 1910
- Synalpheus anasimus Chace, 1972
- Synalpheus anceps Banner, 1956
- Synalpheus ancistrorhynchus De Man, 1909
- Synalpheus androsi Coutière, 1909
- Synalpheus antenor De Man, 1910
- Synalpheus arostris Wicksten, 1989
- Synalpheus bannerorum Abele, 1975
- Synalpheus barahonensis Armstrong, 1949
- Synalpheus belizensis Anker & Tóth, 2008
- Synalpheus bispinosus De Man, 1910
- Synalpheus bituberculatus De Man, 1910
- Synalpheus biunguiculatus
- Synalpheus bocas Anker & Tóth, 2008
- Synalpheus bousfieldi Chace, 1972
- Synalpheus bradleyi Verrill, 1922
- Synalpheus brevicarpus
- Synalpheus brevidactylus Anker & Tóth, 2008
- Synalpheus brevifrons Chace, 1972
- Synalpheus brevispinis Coutière, 1909
- Synalpheus brooksi Coutière, 1909
- Synalpheus carinatus
- Synalpheus carpenteri MacDonald & Duffy, 2006
- Synalpheus chacei Duffy, 1998
- Synalpheus charon
- Synalpheus comatularum
- Synalpheus corallinus MacDonald, Hultgren & Duffy, 2009
- Synalpheus coutierei Banner, 1953
- Synalpheus cretoculatus Banner & Banner, 1979
- Synalpheus crosnieri Banner & Banner, 1983
- Synalpheus curacaoensis Schmitt, 1924
- Synalpheus dardeaui
- Synalpheus demani Borradaile, 1900
- Synalpheus digueti Coutière, 1909
- Synalpheus disparodigitus Armstrong, 1949
- Synalpheus dominicensis Armstrong, 1949
- Synalpheus dorae Bruce, 1988
- Synalpheus duffyi Anker & Tóth, 2008
- Synalpheus echinus Banner & Banner, 1975
- Synalpheus elizabethae
- Synalpheus filidigitus Armstrong, 1949
- Synalpheus fossor
- Synalpheus fritzmuelleri Coutière, 1909
- Synalpheus gambarelloides
- Synalpheus goodei Coutière, 1909
- Synalpheus gracilirostris De Man, 1910
- Synalpheus haddoni Coutière, 1900
- Synalpheus harpagatrus Banner & Banner, 1975
- Synalpheus hastilicrassus Coutière, 1905
- Synalpheus heardi Dardeau, 1984
- Synalpheus hemphilli Coutière, 1909
- Synalpheus herdmaniae Lebour, 1938
- Synalpheus heroni Coutière, 1909
- Synalpheus herricki Coutière, 1909
- Synalpheus hilarulus De Man, 1910
- Synalpheus hoetjesi Hultgren, MacDonald & Duffy, 2010
- Synalpheus idios
- Synalpheus iocasta De Man, 1909
- Synalpheus iphinoe De Man, 1909
- Synalpheus irie MacDonald, Hultgren & Duffy, 2009
- Synalpheus jedanensis De Man, 1909
- Synalpheus kensleyi
- Synalpheus kuadramanus Hultgren, MacDonald & Duffy, 2010
- Synalpheus kusaiensis Kubo, 1940
- Synalpheus laevimanus
- Synalpheus lani Hermoso & Alvarez, 2005
- Synalpheus laticeps Coutière, 1905
- Synalpheus lockingtoni Coutière, 1909
- Synalpheus longicarpus
- Synalpheus lophodactylus Coutière, 1908
- Synalpheus macromanus Edmondson, 1925
- Synalpheus mcclendoni Coutière, 1910
- Synalpheus merospiniger Coutière, 1908
- Synalpheus mexicanus Coutière, 1909
- Synalpheus microneptunus Hultgren, MacDonald & Duffy, 2011
- Synalpheus minus
- Synalpheus modestus De Man, 1909
- Synalpheus mortenseni Banner & Banner, 1985
- Synalpheus mulegensis Ríos, 1992
- Synalpheus mushaensis Coutière, 1908
- Synalpheus neomeris
- Synalpheus nilandensis Coutière, 1905
- Synalpheus nobilii Coutière, 1909
- Synalpheus obtusifrons Chace, 1972
- Synalpheus occidentalis Coutière, 1909
- Synalpheus odontophorus De Man, 1909
- Synalpheus orapilosus Hultgren, MacDonald & Duffy, 2010
- Synalpheus osburni Schmitt, 1933
- Synalpheus otiosus Coutière, 1908
- Synalpheus pachymeris Coutière, 1905
- Synalpheus pandionis Coutière, 1909
- Synalpheus paradoxus Banner & Banner, 1981
- Synalpheus paralaticeps Banner & Banner, 1982
- Synalpheus paraneomeris Coutière, 1905
- Synalpheus paraneptunus Coutière, 1909
- Synalpheus parfaiti
- Synalpheus paulsonoides Coutière, 1909
- Synalpheus pectiniger Coutière, 1907
- Synalpheus peruvianus Rathbun, 1910
- Synalpheus pescadorensis Coutière, 1905
- Synalpheus pinkfloydi De Grave, 2017
- Synalpheus plumosetosus MacDonald, Hultgren & Duffy, 2009
- Synalpheus pococki Coutière, 1898
- Synalpheus quadriarticulatus Banner & Banner, 1975
- Synalpheus quadrispinosus De Man, 1910
- Synalpheus quinquedens Tattersall, 1921
- Synalpheus rathbunae Coutière, 1909
- Synalpheus readi Banner & Banner, 1972
- Synalpheus recessus Abele & W. Kim, 1989
- Synalpheus redactocarpus Banner, 1953
- Synalpheus regalis Duffy, 1996
- Synalpheus riosi Anker & Tóth, 2008
- Synalpheus ruetzleri MacDonald & Duffy, 2006
- Synalpheus sanctithomae Coutière, 1909
- Synalpheus sanjosei Coutière, 1909
- Synalpheus sanlucasi Coutière, 1909
- Synalpheus scaphoceris Coutière, 1910
- Synalpheus sciro Banner & Banner, 1975
- Synalpheus senegambiensis Coutière, 1908
- Synalpheus septemspinosus De Man, 1910
- Synalpheus sladeni Coutière, 1908
- Synalpheus somalia Banner & Banner, 1979
- Synalpheus spinifrons
- Synalpheus spiniger
- Synalpheus spongicola Banner & Banner, 1981
- Synalpheus stimpsonii
- Synalpheus streptodactylus Coutière, 1905
- Synalpheus stylopleuron Hermoso Salazar & Hendrickx, 2006
- Synalpheus superus Abele & W. Kim, 1989
- Synalpheus tenuispina Coutière, 1909
- Synalpheus thai Banner & Banner, 1966
- Synalpheus theano De Man, 1910
- Synalpheus thele MacDonald, Hultgren & Duffy, 2009
- Synalpheus tijou Banner & Banner, 1982
- Synalpheus townsendi Coutière, 1909
- Synalpheus triacanthus De Man, 1910
- Synalpheus tricuspidatus
- Synalpheus tridentulatus
- Synalpheus trispinosus De Man, 1910
- Synalpheus triunguiculatus
- Synalpheus tropidodactylus Banner & Banner, 1975
- Synalpheus tuthilli Banner, 1959
- Synalpheus ul
- Synalpheus wickstenae Hermoso Salazar & Hendrickx, 2006
- Synalpheus williamsi Ríos & Duffy, 1999
- Synalpheus yano