419 
National Marine 
Spencer F. Baird U 
Fisheries Service 
Fishery Bulletin 
First U.S. Commissioner ft- B 
NOAA 
fir established in 1881 
of Fisheries and founder tgJB. 
of Fishery Bulletin 
Abstract— Red grouper ( Epineph - 
elus morio) modify their habitat by 
excavating sediment to expose rocky 
pits, providing structurally complex 
habitat for many fish species. Sur- 
veys conducted with remotely op- 
erated vehicles from 2012 through 
2015 were used to characterize fish 
assemblages associated with grouper 
pits at Pulley Ridge, a mesophotic 
coral ecosystem and habitat area 
of particular concern in the Gulf 
of Mexico, and to examine whether 
invasive species of lionfish ( Pterois 
spp.) have had an effect on these as- 
semblages. Overall, 208 grouper pits 
were examined, and 66 fish species 
were associated with them. Fish as- 
semblages were compared by using 
several factors but were considered 
to be significantly different only on 
the basis of the presence or absence 
of predator species in their pit (no 
predators, lionfish only, red grou- 
per only, or both lionfish and red 
grouper). The data do not indicate a 
negative effect from lionfish. Abun- 
dances of most species were higher 
in grouper pits that had lionfish, 
and species diversity was higher in 
grouper pits with a predator (lion- 
fish, red grouper, or both). These re- 
sults may indicate that grouper pits 
are a favorable habitat for both lion- 
fish and native fish species or that 
the presence of lionfish is too recent 
to have caused changes to fish com- 
munity structure. 
Manuscript submitted 15 August 2016. 
Manuscript accepted 2 June 2017. 
Fish. Bull. 115:419-432 (2017). 
Online publication date: 23 June 2017. 
doi: 10.7755/FB.115.3.11 
The views and opinions expressed or 
implied in this article are those of the 
author (or authors) and do not necessarily 
reflect the position of the National 
Marine Fisheries Service, NOAA. 
Fish assemblages associated with red grouper 
pits at Pulley Ridge, a mesophotic reef in the 
Gulf of Mexico 
Stacey L. Harter (contact author ) 1 
Heather Moe 1 
John K. Reed 2 
Andrew W. David 1 
Email address for contact author: stacey.harter@noaa.gov 
1 Southeast Fisheries Science Center 
National Marine Fisheries Service, NOAA 
3500 Delwood Beach Road 
Panama City, Florida 32408 
2 Harbor Branch Oceanographic Institute 
Florida Atlantic University 
5600 U.S. 1 North 
Fort Pierce, Florida 34946 
The red grouper ( Epinephelus morio ) 
has been harvested in the United 
States since the 1880s and is cur- 
rently the most common grouper spe- 
cies landed in both commercial and 
recreational fisheries of the Gulf of 
Mexico (Fisheries Statistics Division, 
National Marine Fisheries Service, 
Annual Commercial Landing Statis- 
tics, website, and Saltwater Recre- 
ational Data and Statistics, website). 
Like other grouper species, the red 
grouper is a slow growing, late ma- 
turing, relatively stationary, and long 
lived fish (Moe 1 ; Jory and Iversen 2 ). 
Adult red grouper inhabit the deeper 
1 Moe, M. A., Jr. 1969. Biology of the 
red grouper Epinephelus morio (Valen- 
ciennes) from the eastern Gulf of Mex- 
ico. Fla. Dep. Nat. Resour. Mar. Res. 
Lab. Prof. Pap. Ser. 10, 95 p. 
2 Jory, D. E., and E. S. Iversen. 1989. 
Species profiles: life histories and en- 
vironmental requirements of coastal 
fishes and invertebrates (South Flori- 
da) — black, red, and Nassau groupers, 
21 p. U.S. Fish Wildl. Serv. Biol. Rep. 
82(11.110). U.S. Army Corps Eng., TR 
EL-82-4. [Available from website.] 
waters (>70 m) of the shelf edge and 
have been known to modify their 
habitat by excavating sediment to 
expose rocky depressions (or pits) on 
the seafloor in areas where vertical 
relief is not already present (Cole- 
man et ah, 2010). Red grouper use 
these pits as their home territories 
(Scanlon et al., 2005). These exca- 
vations increase the architectural 
complexity of the habitat, attract- 
ing many reef-associated species and 
providing shelter for juveniles of 
some economically important species 
and, thereby, increasing biodiversity 
(Coleman et al., 2010). By excavat- 
ing the sediments, red grouper act as 
“ecological engineers” and may play 
an important role in influencing com- 
munity dynamics (Jones et al., 1994; 
Coleman and Williams, 2002; Cole- 
man et al., 2010). 
Grouper pits have been described 
for 2 marine protected areas (MPAs) 
in the northeastern Gulf of Mexico: 
Madison-Swanson and Steamboat 
Lumps MPAs, which were estab- 
lished in 2000 on the shelf break (at 
