Stevens and Guida: Biological parameters of Chaceon quinquedens in the Mid-Atlantic Bight 
357 
of 33% ovigerous crab. Lack of annual mating for red 
deepsea crab was previously hypothesized by other au- 
thors (Weinberg and Keith, 2003). Female C. maritae in 
South Africa were also reported to have asynchronous 
molting and aseasonal reproductive cycles (Melville- 
Smith, 1989). 
Reproductive cycles >1 year may occur among crabs 
living at extremely low temperatures. Blue king crab 
in the Bering Sea living at temperatures ranging from 
-1°C to 4°C reproduce biennially (Somerton and Ma- 
cintosh, 1985; Jensen and Armstrong, 1989; Stevens 
et al., 2008), and approximately 50% of females bear 
fertilized embryos each spring. Female snow crab re- 
produce biennially in the Gulf of St. Lawrence at tem- 
peratures <1.0°C and have only 2 broods over their 
reproductive lifespan (Moriyasu and Lanteigne, 1998; 
Comeau et al., 1999), whereas snow crab living at high- 
er temperatures (and greater depths) reproduce annu- 
ally, potentially producing 4 lifetime broods (Kuhn and 
Choi, 2011). 
Switching from biennial to annual reproduction can 
potentially halve or double lifetime reproductive output, 
depending on direction of change (Webb et al., 2007). A 
change of only 1°C can advance or delay hatching of 
red king crab (Paralithodes camtschaticus) by about 2 
weeks, possibly contributing to year class failure from 
a mismatch between larval hatching and food sources 
(Stevens et al., 2008). Other crabs with reproductive 
cycles longer than 1 year include the golden king crab 
{Lithodes aequispinus) and others that live at depths 
>500 m and produce lecithotrophic larvae (Shirley and 
Zhou, 1997; Paul and Paul, 2001). Biennial spawning 
has been reported only for crab species living at tem- 
peratures <4°C, and for those with lecithotrophic lar- 
vae, but has not been reported for crabs with plankto- 
trophic larvae living at temperatures >6°C. Therefore, 
crabs of the genus Chaceon are unusual in this respect. 
The extremely low proportion of egg-bearing females 
and the advance stage of egg development observed in 
July 2013 indicate that hatching was nearly completed 
at that time, and some crab bore newly fertilized eggs 
in very early developmental stages. These observations 
indicate that hatching and ovulation events are sepa- 
rated by a time interval, possibly up to a year in red 
deepsea crab (Brachyura). This separation of events is 
in contrast with the reproductive strategy of red king 
crab (Anomura), which molt, mate, and extrude a new 
clutch of eggs within hours after releasing larvae (Ste- 
vens and Swiney, 2007). All species of king crab lack 
the ability to store sperm, however, and, as a result, 
comparisons to brachyurans are more appropriate. 
Snow crab (Brachyura) can store sperm, and multipa- 
rous crab may use stored sperm to produce additional 
clutches of eggs; however, those crab that re-mate must 
do so within 4-7 days after releasing larvae in order 
to produce viable clutches (Paul and Adams, 1984). 
Therefore, it appears that red deepsea crab differ from 
species of Chionoecetes in their ability to separate the 
process of larval release (and presumably mating) from 
that of ovulation and fertilization. 
The current fishery for red deepsea crab is expand- 
ing into the southern portion of the Mid-Atlantic Bight, 
including the Norfolk and Washington Canyon areas. 
This expansion is a result of changing abundance, as 
well as of changes in the availability of processing fa- 
cilities close to fishing grounds (Williams'^). Therefore, 
the population targeted by the fishery is different from 
that fished 20 years or even 10 years previously. The 
results of this study lay a foundation for other syn- 
optic studies on red deepsea crab in the Mid-Atlantic 
Bight. As we went to press (May 2016), we were con- 
tinuing our work with red deepsea crab, analyzing tis- 
sue samples from crab collected during the cruises in 
2011-2013, as well as from samples collected aboard 
commercial fishing vessels in 2014 and 2015. The re- 
sults of this further analysis should reveal more details 
about the biology of the red deepsea crab, including 
size at maturity, reproductive cycles, fecundity, and em- 
bryonic and larval development. 
Acknowledgments 
This research was conducted with partial funding 
from the NOAA Living Marine Resources Cooperative 
Science Center, NOAA grant NA11SEC4810002. We 
gratefully acknowledge the assistance of the officers 
and crews of the NOAA Ships Delaware II and Gordon 
Gunter. We also appreciate the efforts of student par- 
ticipants on all 3 cruises, who were too numerous to 
name individually; still, major assistance to this proj- 
ect was provided by B.-J. Peemoeller, A. Stoneman, and 
I. Suyuheda. We thank R. Langton for his service as co- 
chief scientist in 2013 and to S. Smith and S. Van Sant 
for their participation as watch supervisors during the 
2012 and 2013 cruises, respectively. Comments from L. 
Stehlik and anonymous reviewers helped improve the 
manuscript. 
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