Marine Biology (2023) 170:80

https://doi.org/10.1007/s00227-023-04231-2

ORIGINAL PAPER

Food to go: prey on the web of Octopus maya reveals its diet
Unai Markaida1

Received: 7 February 2023 / Accepted: 5 May 2023

© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2023

Abstract
During an experiment of Octopus maya fishing with baited lines during 2012 and 2013 off Lerma, Campeche, Mexico, 307
octopuses (18.3% of all sampled) shown recently caught 424 prey items in their webs. Preys were composed of crustaceans,
mollusks, teleost fish, and sipunculans representing at least 52 species. Most prey occurrences were single (73.29%), but up
to five prey items were found in some octopus. Crustaceans were the most frequent and abundant prey accounting for 61%FO,
59%N, and 20%W. The most important single prey was the majoid crab Pitho anisodon with 24%FO and 22%N. Molluscans
occurred in half of octopus and represented more than a third of all preys and most of preys’ weight. They were mostly
bivalves. Fishes did not reach 2% of prey share by any account. Prey numbers differed significantly by year. O. maya feed on
small preys, crustaceans, and bivalves in the range of 5–50 mm and gastropod and fishes mostly 50–120 mm in length. Size
of prey was weakly or not related to octopus size, although only larger octopus feed on larger prey such strombid conchs.
This work represents the first systematic study of octopus diet from direct observations on their recently taken prey. The
advantages of this new method to study octopus food are discussed, as well as the feeding behavior associated to some prey.

Keywords Octopus maya · Direct observation · Food · Feeding habits · Prey size

Introduction and McCabe 2014). Gastrointestinal contents, otherwise,

Octopuses are carnivorous predators which feed mainly on
crustaceans, mollusks, and teleost fish. Their food habits
have been study mainly through two methods: examination
of their gut contents and examination of their ‘middens’:
the refuse heaps around their dens (Hanlon and Messenger
2018). However, both methods are largely biased. Items sur-
rounding an octopus den are easily modified by physical
forces such as currents or by other organisms, and this bias
argues against their use in quantitative studies as over-repre-
sents heavy-shelled molluscan share in octopus diet against
crustaceans (Ambrose and Nelson 1983; Ambrose 1984;
Aronson 1989; Mather 1991; Smith 2003). Middens do not
show the whole variety of prey consumed as smaller prey
are ingested completely or feeding occurs away from the den
(Hartwick et al. 1981; Smale and Buchan 1981; Kuhlmann
Responsible Editor: R. Villanueva.
* Unai Markaida
umarkaida@ecosur.mx
1
Laboratorio de Pesquerías Artesanales, El Colegio de
la Frontera Sur (CONACyT), Lerma, 24500 Campeche,
Mexico
underestimate the molluscan component, as hard parts of
crustaceans and teleosts are emphasized. Prey identification
is hard and prey size is also difficult to estimate (Ambrose
and Nelson 1983; Gonçalves and Martins 1994; Dantas et al.
2020; Smith 2003; Ibáñez et al. 2021).
Given the limitations of those indirect methods, Mather
(1991) emphasized the need to use direct observation to
study food choice, foraging behavior, and food handling by
octopuses. Although this seems the simplest and most obvi-
ous technique, it is highly time and effort demanding and
observer’s presence is interfering (Dantas et al. 2020). A few
observations are anecdotal accounts on feeding on unusual
prey such as vertebrates or rare fish (Buxton and Branch
1983; Anderson and Shimek 2014; Muller et al. 2022).
Octopuses carry their recent preys under the arm web while
hunting (Yarnall 1969; Hartwick et al. 1981), and direct
observations on octopus feeding or on food items brought
that way provided limited prey listings (Hochberg and Couch
1971; Smale and Buchan 1981; Aronson 1989; Mather 1991;
Norman 1992; Forsythe and Hanlon 1997; Gilchrist 2003;
Huffard 2007; Silva et al. 2018; Bennice et al. 2021). Only
two studies quantified food items using direct observation.
Armendáriz-Villegas et al. (2014) sampled nine octopus

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which were feeding at the time of their capture, from which
seven had bivalves. Smith (2003) collected 29 prey items
dropped by octopus while fleeing, from which 17 were Hali-
otis midae abalone.
The Mayan octopus Octopus maya is an abundant ben-
thic predator endemic to the shallow waters Campeche Bank
off the Yucatan Peninsula, Mexico. Despite its ecological
importance, its natural diet is poorly known. Seashells from
their middens included the bivalves Codakia orbicularis,
Arca zebra, Modiolus americanus, Trachycardium isocar-
dia, and the gastropod Sinum perspectivum (Solís-Ramírez
1998). The stomach contents of 47 individuals contained
unidentified crustaceans and fish scales (Solís-Ramírez
1997).
Octopus maya is fished with lines baited with dead, fresh
crabs. During a fishing experiment using artificial lures off
Lerma, Campeche (Markaida et al. 2019), a frequent pres-
ence of preys in the web of octopus taken on board was
noticed. Their analysis provided here the first description
using this technique as well as the first methodical account
of O. maya natural prey. Preys were in good shape and pro-
vided new insights in feeding behavior and prey handling
by octopus.
Materials and methods
During the O. maya fishing seasons (August to mid-Decem-
ber) of 2012 and 2013, several fishing trips were made off
Lerma (Campeche) to test for the performance of artificial
lures against natural baits in the fishing lines for octopus
(Markaida et al. 2019). Experiments were performed in shal-
low waters (5–9 m depth) about 0.5–23 km offshore. Fishing
operations typically started at 7 am and ended between 11
am and 1 pm. Lines with both artificial lures and natural
baits caught octopus showing prey in their webs. Stone crab
Menippe mercenaria and warrior swimming crab, Calli-
nectes bellicosus, as well as discarded heads of finfish were
used as natural bait, but they were all easily distinguished
from octopus natural prey. Details on fishing gear and opera-
tion are given elsewhere (Markaida et al. 2019).
All octopuses taken on board were carefully inspected
for prey in their ventral web and arms. They were sacrificed
by professional fishermen with a fast cut between the eyes.
Octopus body weight (BW) was taken to the nearest gram
and measured for mantle length (ML) to the nearest mm.
Sex and maturity were assigned upon examination of repro-
ductive organs following a scale of four maturity stages for
each sex: immature, maturing, mature, and spent (Markaida
et al. 2017).
All prey items were collected and stored in ethanol at
70% until examination. Prey was identified using available
guides for crustaceans, mollusks, and fishes from the Gulf
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Marine Biology (2023) 170:80
of Mexico (Williams 1984; McEachran and Fechhelm 2005;
Tunnell et al. 2010). Alpheid shrimps were additionally col-
lected as reference from commercial crab traps in neighbor-
ing waters of Campeche and identified by Arthur Anker.
Preys were measured to the nearest 0.1 mm for carapace
width in brachyurans, cephalothorax length in other crusta-
ceans, shell length in bivalves and gastropods, and standard
length in fishes. They were weighted to the nearest 0.01 g
for total weight including their shell. Flesh of mollusks
was weighted when available. All prey items were checked
for drill and bite marks in crustacean carapaces, eyes, and
appendages, and in mollusc shells (Dodge and Scheel 1999;
Pech-Puch et al. 2016).
Diet quantification was given as frequency of occurrence
(FO), number (N), weight (W), and their respective percent-
ages (%FO, %N, and %W) (Urrutia-Olvera et al. 2021). Prey
numbers (%N) were used to test for significant dietary differ-
ences between years through an ANOSIM (PRIMER 6.1.6).
Preys were grouped into major taxa: Majoid crabs, other
brachyurans, other crustaceans, bivalves, gastropods, fish,
and others. Octopus samples were grouped daily, accounting
for 31 days with samples in 2012 versus 39 days in 2013.
Data were transformed using the square root and analyzed by
the Bray–Curtis similarity measure (Smith 2003). A Simper
test was performed to determine which prey groups most
contributed to the total average dissimilarity.
To test the fact that prey diversity increases with octopus
size (Smale and Buchan 1981; Smith 2003), a regression was
performed for prey species number by each octopus 100 g
BW interval. Regressions of prey size on octopus weight
were run to determine any relation.
Results
During the 2012 season, 391 O. maya were sampled
for 33 days (26 September to 13 December) from which
90 (23%) had preys in their webs. In 2013, 40 days were
sampled (2 August to 4 October) for 1,283 octopus from
which 217 (16.9%) yielded preys. Overall, prey-bearing
octopus accounted for 307 or 18.3% of all octopuses sam-
pled. They were 125 females, 169 males and 13 unsexed
octopus. Females were mainly immature while most males
were mature, as expected during the fishing season (Fig. 1;
Markaida et al. 2017). They averaged 698 ± 322 g in BW,
ranging 100–1940 g BW, and they did not differ in BW by
sex (Mann–Whitney U test, U = 9525, N1 = 125, N2 = 169,
P > 0.05).
A total of 424 prey items (127 in 2012 and 297 in 2013)
were found in the webs of these 307 octopus. Most prey
occurrences were single (73.29%), 19.87% were double,
3.26% triple, 2.93% quadruple and only two octopus (0.65%)
had five prey items.
Marine Biology (2023) 170:80
Fig. 1  Cumulative frequency
distribution of the body weight
of Octopus maya which show
preys in their webs: a females
and b males, by maturity stage
At least 52 species of preys were found (Table 1). They
belong to crustaceans, mollusks, teleost fish, and sipun-
culans. Crustaceans were the most frequent and abundant
of such groups, accounting for 61%FO and 59%N. In this
group, brachyurans accounted for 53%FO and 51%N and
the most important single prey was the majoid crab Pitho
anisodon, with 24%FO and 22%N. Molluscans occurred in
half of octopus (47%FO) and represented more than a third
of all preys (38%FO) and most of preys’ weight (78%W).
They were mostly bivalves (36%FO and 29%N). Fishes did
not reach 2% of prey share by any account (Table 1).
Prey numbers differed significantly by year, although
the R value was quite low (ANOSIM: R = 0.145, P < 0.05).
Simper analysis determined that majoid crabs (23.27%),
other brachyurans (22.52%) and bivalves (21.71%) contrib-
uted most to the total average dissimilarity. Despite similar
numbers for yearly brachyuran share in octopus webs, P.
anisodon participation by %FO and %N halved from 2012
to 2013, while that of the rest of brachyurans doubled. Gas-
tropods’ share in the diet also more than doubled from 2012
to 2013. The rest of the prey did not change in the share
(Table 1). Prey diversity (species number) for both years
pooled was not related to octopus size (r2 = 0.002, P > 0.05),
but it was strongly correlated with sample size in octopus
size intervals (r2 = 0.84, P < 0.01).
Octopus maya feed on small preys. Crustaceans and
bivalves were in the range of 5–10 to 45–50 mm. Less-abun-
dant gastropods and fishes were mostly 50 to 100–120 m in
length (Table 2, Fig. 2). P. anisodon ranged 7.3–25.2 mm
CW. They were 38 females and 37 males, and they did not
differ significantly for CW (t = − 0.57, df = 73; P > 0.05).
Ten P. anisodon females 15.8–19.7 mm CW had eggs.
Smaller preys were represented by alpheids and xanthoid,
and Pilumnus crabs < 10 mm. The largest preys were Mac-
rostrombus costatus conchs, which accounted for most of
the weight (Table 2).
The ghost shrimp Glypturus acanthochirus and the
giant hermit crab Petrochirus diogenes were represented
just by one of their chelipeds. The former was represented
by three major and two minor chelipeds. A single case
Page 3 of 12 80
of cannibalism was represented by a 700 g BW female
carrying a 10 g BW O. maya lacking five arms. Multiple
occurrences of four and five preys were multi-specific,
involving crustaceans and mollusks, except for one octo-
pus bringing five Strombus pugilis and another carrying
four P. anisodon. Those five S. pugilis were 44–50 mm in
length, intact, and with no drills. All other occurrences of
this prey were single.
Size of P. anisodon was weakly related to octopus size
(r2 = 0.13; P < 0.001), as well as size of other brachyurans
(r2 = 0.09; P < 0.01). Size of bivalves and S. pugilis did not
relate to octopus size (P > 0.05). Size of gastropods exclud-
ing S. pugilis did very significantly relate to octopus size
(r2 = 0.63; P < 0.01), mainly because largest prey (≥ 90 mm
length) M. costatus and Fasciolaria tulipa were taken by
large octopus > 950 g BW (Fig. 3). Strombid conchs repre-
sented over half of total prey weight (Table 1).
Most preys in octopus web were in good condition, with
most crustaceans complete or just lacking some append-
ages and most mollusks intact (Table 3). This fact allowed
to identify 99.9% of all preys. Unidentified preys (Table 1)
were mainly lost samples or items eaten beyond recognition.
Crustacean exoskeletons were very rarely drilled. Carapaces
were drilled in two cases. A Mithraculus forceps had many
drill attempts and a blue crab Callinectes sp. had three drills.
All other drills were made on the palms of different species.
Three out of the five Glypturus palms had an incomplete
drill each (Fig. 4). Bites were found in carapaces and palms
of three species of crabs. Punctured eyes were observed in
three Pitho and two Speocarcinus crabs (Table 3).
Two species of bivalves with thin shells, Tagelus and
Modiolus, showed sometimes broken valves. From 25 Tage-
lus represented by both complete valves, only four shells had
all the hinge teeth. Four more had three teeth left, 11 had two
teeth each, nine had only one tooth and in four all teeth were
gone. Drilled holes were only found in three bivalves with
thick shell, a Chione elevata and two Megapitaria maculata.
Their drill holes were always located in the valve periphery
(Table 3; Fig. 4). Roughly half of thin-shelled and most of
thick-shelled bivalves had flesh remains.
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 80 Page 4 of 12 Marine Biology (2023) 170:80

Table 1  Summary of preys 2012 2013

found in the web of Octopus
maya FO %FO N %N W (g) %W FO %FO N %N W (g) %W

CRUSTACEA 59 65.5 79 62.2 175.2 19.7 130 59.9 173 58.2 229.4 21.0
BRACHYURA​ 48 53.3 66 52.0 134.9 15.2 116 53.5 150 50.5 212.9 19.5
Pitho anisodon 33 36.7 43 33.9 65.9 7.4 42 19.4 51 17.2 70.4 6.4
Pitho lherminieri 1 0.5 1 0.3 0.9 0.1
Mithraculus forceps 2 2.2 4 3.1 5.3 0.6 4 1.8 4 1.3 3.7 0.3
Nonala holderi 1 1.1 1 0.8 4.2 0.5 1 0.5 1 0.3 0.6 0.1
Macrocoeloma trispinosum 2 2.2 2 1.6 6.6 0.7
Hepatus epheliticus 4 4.4 4 3.1 21.6 2.4 10 4.6 12 4.0 29.6 2.7
Calappa sulcata 1 1.1 1 0.8 9.9 1.1 3 1.4 3 1.0 16.6 1.5
Callinectes spp. 13 6.0 13 4.4 9.5 0.9
Portunus sayi 3 3.3 3 2.4 8.2 0.9 5 2.3 4 1.3 2.9 0.3
Unidentified Portunidae 1 1.1 1 0.8 0.0 1 0.5 1 0.3 0.07 0.0
Persephona mediterranea 3 3.3 3 2.4 6.2 0.7 15 6.9 15 5.1 18.4 1.7
Acanthilia intermedia 1 0.5 1 0.3 4 0.4
Euryplax nitida 1 1.1 1 0.8 0.5 0.1 13 6.0 16 5.4 6.87 0.6
Speocarcinus carolinensis 2 2.2 2 1.6 6.3 0.7 4 1.8 5 1.7 20.74 1.9
Panoplax depressa 1 0.5 1 0.3 0.1 0
Micropanope sculptipes 1 0.5 1 0.3 0.02 0
Panopeus herbstii 1 0.5 1 0.3 0.4 0
Platylambrus serratus 10 4.6 12 4.0 26.7 2.4
Pilumnus sayi 1 0.5 1 0.3 1.2 0.1
Pilumnus sp. 1 0.5 1 0.3 0.06 0
Pinnixa sp. 1 0.5 1 0.3 0.03 0
Unidentified Brachyura 1 1.1 1 0.8 4 1.8 4 1.3
OTHER CRUSTACEA 13 14.4 13 10.2 40.3 4.6 21 9.7 23 7.7 16.5 1.5
Alpheus mathewsae 6 2.8 7 2.4 2.1 0.2
Alpheus cf. packardii 1 1.1 1 0.8 0.1 0.0 3 1.4 4 1.3 0.2 0
Unidentified Alpheus 3 3.3 3 2.4 0.7 0.1 1 0.5 1 0.3 0.1 0
Penaeus duorarum 3 3.3 3 2.4 8.6 1.0 3 1.4 3 1.0 5.9 0.5
Rimapenaeus constrictus 1 0.5 1 0.3 0.4 0
Glypturus acanthochirus 1 1.1 1 0.8 2.4 0.3 4 1.8 4 1.3 7 0.6
Paraxiopsis gracilimana 1 0.5 1 0.3 0.05 0
Paguristes puncticeps 1 1.1 1 0.8 1.3 0.1 1 0.5 1 0.3 0.1 0
Petrochirus diogenes 1 1.1 1 0.8 24.9 2.8
Gibbesia prasinolineata 1 1.1 1 0.8 2.5 0.3
Unid. Crustacea 2 2.2 2 1.6 1 0.5 1 0.3 0.5 0.1
MOLLUSCA 38 42.2 46 36.2 694.1 78.2 106 48.8 117 39.4 849.3 77.7
BIVALVIA 33 36.7 41 32.3 93.5 10.5 77 35.5 83 27.9 207 19
Tagelus divisus 7 7.8 7 5.5 2.9 0.3 27 12.4 28 9.4 12.9 1.2
Laevicardium serratum 14 15.6 15 11.8 35.1 4.0 17 7.8 17 5.7 51.3 4.7
Dallocardia muricata 3 1.4 3 1.0 2.1 0.2
Chione elevata 4 4.4 5 3.9 21.3 2.4 12 5.5 12 4.0 66.3 6.1
Megapitaria maculata 2 2.2 2 1.6 20.4 2.3 7 3.2 7 2.4 46.2 4.2
Dosinia elegans 1 1.1 1 0.8 1.6 0.2 4 1.8 6 2.0 15.9 1.5
Modiolus squamosus 6 6.7 7 5.5 12.2 1.4 5 2.3 5 1.7 7.7 0.7
Mactrotoma fragilis 2 0.9 2 0.7 2.2 0.2
Anatina anatina 1 0.5 1 0.3 2.3 0.2
Unid. Bivalvia 4 4.4 4 3.1 2 0.9 2 0.7
GASTROPODA 5 5.6 5 3.9 600.6 67.7 28 12.9 33 11.1 632.1 57.9

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Marine Biology (2023) 170:80 Page 5 of 12 80

Table 1  (continued) 2012 2013

FO %FO N %N W (g) %W FO %FO N %N W (g) %W

Strombus pugilis 21 9.7 25 8.4 552.9 50.6

Macrostrombus costatus 2 2.2 2 1.6 568 64 1 0.5 1 0.3
Prunum labiatum 1 1.1 1 0.8 5.53 0.6 3 1.4 3 1.0 14.2 1.3
Fasciolaria tulipa 1 1.1 1 0.8 26.9 3.0 1 0.5 1 0.3 50.1 4.6
Oliva sayana 1 0.5 2 0.7 8.6 0.8
Naticarius canrena 1 0.5 1 0.3 6.2 0.6
Crepidula convexa 1 1.1 1 0.8 0.21 0
Octopus maya 1 0.5 1 0.3 10.1 0.9
PISCES 2 2.2 2 1.6 17.57 2.0 4 1.8 4 1.3 11.2 1.0
Lutjanus synagris 2 0.9 2 0.7 10.3 0.9
Orthopristis chrysoptera 1 1.1 1 0.8 11 1.2
Paraclinus cingulatus 1 0.5 1 0.3 0.2 0
Opistognathus sp. 1 1.1 1 0.8 6.57 0.7
Unid. Teleost 1 0.5 1 0.3 0.7 0.1
Sipuncula 3 1.4 3 1.0 1.9 0.2
TOTAL 90 100 127 100 886.9 100 217 100 297 100 1091.8 100

Diet quantification is given per year as frequency of occurrence (FO), number (N) and weight (W, in
grams), and their respective percentages (%FO, %N and %W)

Shells of most gastropods species and individuals were
drilled. All drilled gastropods were otherwise intact (includ-
ing operculum). Drill holes were located on the spire (only
twice occurred on the body whorl) and mostly on the aper-
ture (ventral) side of the shell; only 5 incomplete drills were
on the dorsal side (Fig. 4). Strombus pugilis showed all
stages of predation. Ten conchs were intact, 12 had shells
drilled and three individuals were lacking the shell (Table 3).
Eight shells had a single complete drill hole, two had a sin-
gle incomplete drill, another had both a complete and an
incomplete drill, and a third conch had three incomplete
drills. Each Fasciolaria shell had three drill holes, and in
one shell, two of them were complete (Fig. 4).
Discussion
This work represents the first systematic study of octopus
diet from direct observations on their recently taken prey.
Catching octopus with baited lines allows the collection of
the prey that the octopus just hunted. It is a friendly and
non-invasive sampling as octopus voluntarily attack and
holds the bait and they are taken on board without damage.
Octopus can be sampled for size and sex and collect the prey
they carry. This method allows detailed identification of both
predator and prey, as most preys were in good condition.
This technique could be routinely used in the few octopus
fisheries, which used baited lines or lures (Sauer et al. 2021).
It could be used also as scientific sampling to study octo-
pus diet in many other places. Although in this experiment
octopuses were sacrificed by fishermen, they could have
been safely released back to the sea. Octopuses sampled
while diving with hooks (Armendáriz-Villegas et al. 2014),
however, are injured in such a way that they more likely
release their prey, rarely retaining items in their web.
However, the method is time-consuming as it took 77
mornings at sea to complete the collection here described.
It provides a clue of instantaneous feeding at the time of
capture, so results could also be different if observed at other
time of the day such as in the afternoon or at night (Smith
2003). Still, prey numbers per octopus are limited to one in
most cases (73%), so precludes the determination of the indi-
vidual specialization degree as commonly done in midden
studies (Anderson et al 2008; Kuhlmann and McCabe 2014).
As such, the number of prey species is strongly related to
octopus sample size. The other direct technique of observ-
ing octopus feeding underwater is in the same way difficult
and time-consuming (Dantas et al. 2020). It provides excel-
lent evidence on octopus’s foraging behavior, but in turn
identification of prey is often tentative (“probable prey” in
Mather 1991).
This method is not as strongly biased toward some prey
groups as they are gastric contents and midden analy-
sis (Dantas et al. 2020), but still has some partialities.
Handling time for different prey items by octopus varies
greatly, so determining the chances to observe them in the
octopus web. Crabs, shrimp, and thin-shelled bivalves are
handled and consumed within a few minutes, while drill-
ing hard-shelled bivalves can take up to hours (Ambrose
1984; Mather 1991; Gilchrist 2003). So larger preys and

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 80 Page 6 of 12 Marine Biology (2023) 170:80

Table 2  Size, relative size to Prey N Size, mm % ML Weight, g

octopus mantle length (% ML)
and mean weight (flesh weight) Mean ± SD Range Mean ± SD Range (Flesh)
of 376 preys of Octopus maya
from Lerma, Campeche BRACHYURA​ 188 19.5 ± 8 5.5–53.4 17 ± 7 4–41 1.7
Pitho anisodon 85 17.2 ± 3.5 7.3–25.2 15 ± 3 8–25 1.4
Pitho lherminieri 1 13.7 17 0.9
Mithraculus forceps 8 16.5 ± 4.1 11–21.5 14 ± 4 10–20 1.1
Nonala holderi 2 18.7 ± 10.3 11.4–26 16 ± 10 8–23 2.4
Macrocoeloma trispinosum 2 28 ± 0.7 27.5–28.5 24 ± 0.3 24–25 3.3
Hepatus epheliticus 13 28.7 ± 9.3 16.5–45.5 25 ± 8 13–41 3.4
Calappa sulcata 3 38.2 ± 2.6 35.8–41 32 ± 5 28–38 6.6
Callinectes spp. 13 28 ± 9.3 11–45.8 24 ± 7 9–31 0.7
Portunus sayi 8 29.7 ± 12.1 16–53.4 27 ± 8 14–36 1.4
Persephona mediterranea 18 17 ± 4.6 11.3–28 15 ± 4 8–22 1.3
Acanthilia intermedia 1 23 18 4
Euryplax nitida 14 11.4 ± 4 6.4–21.7 10 ± 3 4–17 0.4
Speocarcinus carolinensis 5 24.4 ± 4.6 20–31.3 20 ± 6 11–27 3.9
Panoplax depressa 1 8.2 0.1
Micropanope sculptipes 1 6.2 8 0.02
Panopeus herbstii 1 10 10 0.39
Platylambrus serratus 9 24.5 ± 4.3 17.6–19 20 ± 5 12–19 2.4
Pilumnus sayi 1 11.5 1.25
Pilumnus sp. 1 6.8 5.1 0.06
Pinnixa sp. 1 5.5 5 1.4
OTHER CRUSTACEA
Alpheus mathewsae 5 9.5 ± 1.7 7–11.2 7±2 5–10 0.3
Alpheus packardii 4 5.3 ± 0.9 4–6.1 4±1 3–5 0.06
Unidentified Alpheus 2 8.2 ± 1 7.5–9 7 ± 0.7 6–7 0.2
Penaeus duorarum 6 15.4 ± 4.4 10.4–21 13 ± 4 7–18 2.4
Rimapenaeus constrictus 1 18.5 15 0.42
Glypturus acanthochirusa 5 24.9 ± 8.5 13.5–33.6 22 ± 9 10–32 1.86
Paguristes puncticeps 2 5 ± 2.7 3.1–7 5±2 3–6 1.1 (0.1)
Petrochirus diogenesa 1 56 41 25
Gibbesia prasinolineata 1 15 14 2.5
BIVALVIA 117 27 ± 6.6 11–45.4 25 ± 8 9–47 2.6(0.7)
Tagelus divisus 35 27.2 ± 2.9 21.6–31.6 25 ± 4 17–38 0.4 (0.2)
Laevicardium serratum 32 23.7 ± 6.2 12.5–36 20 ± 6 10–40 2.7 (0.8)
Dallocardia muricata 3 16.9 ± 7.4 11–25.2 16 ± 10 9–58 0.7
Chione elevata 17 27 ± 2.4 23.7–30.5 26 ± 6 19–36 5.1 (0.5)
Megapitaria maculata 9 35 ± 7.4 24.1–44.4 33 ± 8 22–47 7.4 (1.4)
Dosinia concentrica 7 26.1 ± 7.5 16.7–36.5 23 ± 8 11–33 2.5 (0.7)
Modiolus squamosus 12 31.1 ± 9.6 17.5–45.4 28 ± 11 12–45 1.8 (0.7)
Mactrotoma fragilis 1 30 30 1.1 (0.2)
Anatina anatina 1 39.3 41 2.3 (0.9)
GASTROPODA 34 60 ± 31.1 13.2–180 53 ± 22 13–119 20.8 (4.4)
Strombus pugilis 22 61 ± 12.7 44.1–79.9 55 ± 11 39–76 25 (3.9)
Macrostrombus costatus 2 143.5 ± 51.6 107–180 97 ± 31 75–119 No record
Prunum labiatum 4 28.2 ± 1 27.2–29.3 28 ± 4 24–32 5 (1.5)
Fasciolaria tulipa 2 97.4 ± 11.5 89.3–105.6 88 ± 6 83–92 38.5 (18.3)
Oliva sayana 2 31.9 ± 2.3 30.3–33.6 31 ± 2 29–32 4.3 (0.8)
Naticarius canrena 1 25.5 20 6.2
Crepidula convexa 1 13.2 13 0.2

13
Marine Biology (2023) 170:80 Page 7 of 12 80

Table 2  (continued) Prey N Size, mm % ML Weight, g

Mean ± SD Range Mean ± SD Range (Flesh)

Octopus maya 1 30 25 10.8

TELEOSTEI 6 91.1 ± 40.9 29–128 71 ± 25 27–100 4.8
Lutjanus synagrisb 2 93 ± 49.4 58–128 71 ± 10 64–78 5.1
Orthopristis chrysopterab 1 125.8 92 11
Paraclinus cingulatus 1 29 27 0.2
Opistognathus sp. 1 86 69 6.6
Unidentified Teleost 1 120 100 0.7
Sipuncula 3 50.3 ± 6.5 44–57 43 ± 8 37–53 0.6

Prey size is given as carapace width in Brachyura, carapace length in other crustaceans, shell length in
Bivalvia and Gastropoda, mantle length in octopus, and standard length in teleosts
a
Prey represented by a cheliped only, Chela length (mm)
b
Standard lengths estimated from fish parts

Fig. 2  Cumulative frequency

distribution of the size of preys
found in the web of Octopus
maya from Campeche: a Pitho
anisodon by crab sex, b Rest of
Brachyurans, c Bivalves and d
Gastropods

hard-shelled mollusks, such as bivalves and gastropods,
could be overrepresented in octopus webs as they take
longer time to handle and consume than smaller and
softer prey. Octopuses busy in consuming large prey also
could not be so interested in the bait. Some of the heavi-
est preys may drop off the webs when the octopus is lifted
by the line, although the shallow waters where O. maya
is fished minimize the time required to pull the octopus
on board, favoring prey retention. On the other side, most
crabs ≤ 10 mm CW were eaten whole by O. vulgaris type
III from South Africa (Smale and Buchan 1981; Smith
2003), so smaller preys, such as alpheid shrimps and xan-
thoid, and pilumnid crabs might be underrepresented in
O. maya webs as they demand short handling time, being
rapidly devoured (Smith 2003; Kuhlmann and McCabe
2014). Future work on O. maya natural feeding should
be complemented with the analysis of stomach contents
and middens, together with more recent methodologies

13
 80 Page 8 of 12
Fig. 3  Relationship between prey size and Octopus maya body
weight: a Brachyurans, solid line is the regression for Pitho anisodon,
dashed line for other brachyurans, b Bivalves and c Gastropods, solid
line is the regression for other gastropods
such as molecular prey identification and stable isotopes
as well (Dantas et al. 2020; Bo et al. 2020; Urrutia-Olvera
et al. 2021).
Octopus maya feed more on crustaceans than on mol-
lusks in Lerma, Campeche. O. insularis (Leite et al. 2009,
2016; Urrutia-Olvera et al. 2021) and “O. vulgaris” from
Honduras (Gilchrist 2003) eat also more crabs than mol-
luscans. In all other studies from the western Atlantic so far
mollusks, mainly bivalves, dominated the diet (Hochberg
and Couch 1971; Aronson 1989; Mather 1991; Gilchrist
13
Marine Biology (2023) 170:80
2003; Anderson et al 2008; Kuhlmann and McCabe 2014;
Leite et al. 2016; Bennice et al. 2021). The dietary differ-
ence between years found off Lerma is very probably due to
fishermen visiting different fishing spots with different prey
availability. This suggests the opportunistic nature of octo-
pus with diet reflecting prey availability at the different sites
and habitats (Smith 2003; Leite et al. 2016). Thus, diet of O.
maya from other localities might reveal variations as well.
The only commercial species seized by octopus were
juvenile specimens of pink shrimp Penaeus duorarum, blue
crab Callinectes, and adult strombid conchs and tulip shells.
No stone crab Menippe mercenaria was found despite being
the commercially most important crab in Campeche–Lerma
area. Although octopuses interfere with some crab and lob-
ster fisheries (Smith 2003; Boyle and Rodhouse 2005), com-
mercial invertebrates are largely absent as natural prey of
octopus from the western Atlantic, except for few portunid
crabs and strombid conchs which are occasionally enlisted
without being important (Hochberg and Couch 1971; Gil-
christ 2003; Anderson et al. 2008; Kuhlmann and McCabe
2014; Bennice et al. 2021; Urrutia-Olvera et al. 2021).
Octopus maya feed on mainly small-sized crabs and
bivalves < 40 mm in length. Their prey size is in the range of
the prey of most octopus species as O. insularis (Leite et al.
2009, 2016; Silva et al. 2018), O. cyanea (Mather 2011), and
O. vulgaris from northeastern Atlantic (Ambrose and Nelson
1983; Gonçalves and Martins 1994), although those from
Vigo eat somewhat larger prey (Guerra and Nixon 1987).
Feeding on small-sized prey coupled with lack of or little
relation with octopus size, suggests that O. maya time-min-
imize their duration of foraging as does O. insularis from
Brazil (Leite et al. 2009, 2016; Silva et al. 2018). Although
O. maya will readily attack large crabs set as bait in the fish-
ing lines (30–80 mm CW for the stone crab Menippe mer-
cenaria; Markaida et al. 2019), their natural preys are much
smaller. Only the largest portunid and calappid crabs found
in octopus web may overlap with the size of smaller baits.
The giant octopus Enteroctopus dofleini from Northwest
Pacific feed on larger prey (modes at 40–90 mm length)
than O. maya. Relationship between E. dofleini size and
prey size in the middens is controversial. Hartwick et al.
(1981) found a weak relation with the size of the clam Cli-
nocardium, while Vincent et al. (1998) did not find it for the
crab Telmessus. In other studies (Dodge and Scheel 1999;
Scheel et al. 2007), however and despite the size of the giant
octopus, most preys (< 30 mm mean lengths) did fall within
prey size taken by O. maya.
E. magnificus from Namibia feed on larger crabs
80–120 mm length, while fishes and hagfish were only
consumed by larger octopus > 1 kg BW (Villanueva 1993).
O. vulgaris type III from South Africa also consume larger
prey (50–80 mm mean lengths) than O. maya. The larg-
est octopus eat the largest mussels and abalone (Smale
Marine Biology (2023) 170:80 Page 9 of 12 80

Table 3  Condition of most abundant prey (numbers) found in the web of Octopus maya
Prey species N Prey stage Chip Drilling Drill ⌀ (mm)
I A C L N% S M In Short × long

Pitho anisodon 94 25 48 11 10 6
Mithraculus forceps 8 3 1 4 12 1 0.6–1.1 × 0.7–1.1
Hepathus ephiliticus 16 9 5 1 1
Calappa sulcata 4 2 1 1
Callinectes spp. 13 4 7 2 7 1 0.3–0.7 × 0.4–0.8
Portunus sayi 8 2 4 2
Persephona 18 13 2 3
Euryplax nitida 17 8 5 1 3 5 2 0.6–1.8 × 0.7–2.0
Speocarcinus 7 4 1 2
Platylambrus 12 7 1 1 3 6 8 1 0.7–0.8 × 1.0–1.3
Alpheus spp. 16 3 5 3 5
Glypturus 5 5 1 60 3 0.4–0.7 × 0.4–0.9
Paraxiopsis 1 1 100 1 1.0 × 0.9
TOTAL 219 80 80 28 31 18 1 3 5
I V B S

Tagelus divisus 35 16 7 7 5
Laevicardium 32 6 3
Chione elevata 17 13 4 1 6 1 0.5–0.9 × 0.7–1.0
Megapitaria 9 8 1 22 2 0.6–0.7 × 0.6–0.8
Dosinia elegans 7 5 2 1 14
Modiolus 12 5 6 4 1
Strombus pugilis 25 22 3 55 8 2 2 0.6–1.3 × 0.7–1.5
Prunum labiatum 4 4 75 3 0.7–0.8 × 0.7–1.1
Fasciolaria tulipa 2 2 100 1 1 0.6–1.0 × 0.8–1.1
Oliva sayana 2 2 50 1 0.7 × 0.9
TOTAL 145 77 13 11 22 5 12 3 6

Crustaceans: I, intact; A, appendages lacking; C, just carapace and L, lacking carapace. Bivalves: I, intact; V, just valves; B, broken valve, S, sin-
gle valve. Number of prey drilled: S, single drill; M, multiple; In, Incomplete. Range of both axis of external orifices in drilled holes

and Buchan 1981; Smith 2003). In the same way, only the
largest O. maya were able to consume the largest prey,
although there is no evidence that they show a wider vari-
ety of prey as seen in South African octopus (Smale and
Buchan 1981; Smith 2003).
Octopus maya do not drill brachyuran carapaces, as
shown in laboratory experiments by Pech-Puch et al.
(2016). They found that octopus saliva is driven through
punctures on the eye or, more often, on the arthrodial
membrane. This fact would explain the high numbers
of lacking appendages in crabs found in octopus webs.
Drill attempts were found in nine crustaceans, but bites
were more numerous. Drill attempts on Glypturus palms
are particularly significant as they are strongly calcified
pieces, as the thick shells of mollusks. Eye punctures were
observed in five crabs, although the preservation stage of
the samples might have precluded observing more.
Octopuses had three penetration techniques in bivalves:
they could pull the valves apart, chip on the valve edge or
drill through the shell (Anderson and Mather 2007). Most
bivalve preys of O. maya, such as Tagelus divisus, Laevicar-
dium serratum and Modiolus squamosus, have thin valves
and they are easily pulled apart or broken. Hinge teeth from
T. divisus shells were usually broken after such forcing.
Bivalves that showed drilled holes, such as Chione elevata
and Megapitaria maculata, have thicker shells. Similarly,
Enteroctopus dofleini and O. mimus easily open or break the
shells of Tagelus and Mytilus, while they must laboriously
drill or chip the shell of the clam Protothaca (Cortez et al.
1998; Anderson and Mather 2007). They concluded that this
differential handling makes octopus to take effort and pro-
cessing time into consideration when choosing prey. Limited
observations suggest that O. maya drill in the valve periph-
ery, the thinnest part of the shell, hence minimizing handling

13
 80 Page 10 of 12
Fig. 4  Location of drill holes made by Octopus maya in a 12 shells
of ventral and dorsal sides of Strombus pugilis, b Two shells of Fas-
ciolaria tulipa; two incomplete drill holes in dorsal side not shown,
c Three shells of Prunum labiatum plus an Oliva sayana in blue, d
Left valve of Chione elevata showing a complete drill hole and a
time as in O. vulgaris from the Mediterranean (Ambrose
and Nelson 1983) and O. dierythraeus (Steer and Semmens
2003), unlike near the myostracum of the abductor muscles
as does O. mimus (Cortez et al. 1998).
Most gastropod species taken by O. maya had drilled
holes in their shells, and possibly all of them would be
drilled as shown in captivity (Pech-Puch et al. 2016). Drilled
holes were made mainly on the ventral side of the spire.
Other octopus species also drilled the spire of gastropods,
although several explanations had been given and reasons
remained unclear (Fee et al. 2023).
Sublethal predation by O. maya was noticed by the sei-
zure of the sole chelipeds of ghost shrimps Glypturus and
the giant hermit crab Petrochirus. Glypturus live in deep
burrows, and O. maya could possibly only harvest one of
their spread chelipeds at the burrow opening. The giant her-
mit crab Petrochirus diogenes uses the chela used to block
the entrance of the shell.
Most preys of O. maya are characteristic of shallow
soft bottoms (Williams 1984; Tunnell et al. 2010). Pitho
anisodon is associated to beds of Thalassia testudinum
(Heck 1977; Sánchez and Raz-Guzman 1997). The genus
Pitho has been found as prey in dens of “O. vulgaris”
and O. insularis from the shallow tropical Western Atlan-
tic, often associated with seagrass beds (Aronson 1989;
Gilchrist 2003; Anderson et al. 2008; Leite et al. 2009;
Kuhlmann and McCabe 2014), contrasting with octopus
from rocky reefs which feed more on other mithracid crabs
(Mithraculus, Mithrax) and xanthids (Mather 1991; Gil-
christ 2003; Anderson et al. 2008; Leite et al. 2009, 2016;
Urrutia-Olvera et al. 2021). Chione elevata is the most
13
Marine Biology (2023) 170:80
chip (arrow), and incomplete drill holes from two Macrollista macu-
lata, and e Chela with incomplete drill holes from three Glypturus
acanthochirus (detail in arrow). White and red/blue circles represent
incomplete and complete drill holes, respectively
common infaunal bivalve in several localities and octo-
pus middens from the western Atlantic (Aronson 1989;
Kuhlmann and McCabe 2014; Silva et al. 2018; Bennice
et al. 2021).
The stage of prey allows to infer on octopus foraging
habits. The octopus found with five intact Strombus pugi-
lis conchs in the web is probably a specialized individual
(Mather 1991). Octopuses preform long foraging trips in
which they visit resting and feeding sites away from dens,
where those conchs were probably collected (Yarnall 1969;
Hochberg and Couch 1971). Octopuses consumed 70% of
their prey at home and 30% away from it (Hartwick et al.
1981; Mather 1991), but as drilling all those conchs would
take many hours, they were most likely being brought to be
consumed safely at the den.
Commercial fishing for octopus in the morning
(Markaida et al. 2019) and laboratory research with juve-
niles (González-Navarrete et al. 2022) suggest that O. maya
is diurnal. Pitho was taken during the day by octopus from
Bahamas (Aronson 1989) using visual detection followed
by a direct visual attack (Forsythe and Hanlon 1997). How-
ever, all bivalves taken by O. maya are shallow infaunal,
including the most common Tagelus which is deep infaunal
and excluding the mussel Modiolus squamosus which is an
epibyssate (Tunnell et al. 2010). In the same way, most of the
diurnal O. cyanea’s potential preys are well hidden during
daylight hours. The main strategy in such case would be to
forage in a tactile feeding and speculative hunting way, such
as inserting the arms into holes in the sand (Forsythe and
Hanlon 1997; Hanlon and Messenger 2018). Unsuccessful
attacks of octopus on fishes have been reported at day (Leite
Marine Biology (2023) 170:80
et al. 2009), although they could have been taken at night
as observations on other octopus suggest (Aronson 1989).
Acknowledgements I am greatly indebted to Rubén Sánchez and octo-
pus fishermen Francisco Deib “Rana”, Alex Jah, and Ramón Méndez
and his sons for their collaboration during the fishing experiment.
Almendra Rodríguez and Ivan Méndez also assisted in the sampling.
José Luis Mier “Sapo” collected the alpheids. Sara Balán and Anabel
León, BIOMARCCA, ECOSUR, Campeche, assisted with the identifi-
cation of some crustaceans. Art Anker, Universidade Federal de Goiás,
Brazil, kindly identified Alpheus mathewsae and A. cf. packardii from
Campeche. The ms greatly benefited from the critical review of two
anonymous reviewers.
Author contributions UM contributed to the study conception and
design, samples collection and examination, data analysis and writing
all parts of the MS including figures.
Funding Fundación Produce Campeche A.C. supported the octopus
experiment through the project “Validation of an alternative to Mayan
octopus fishing (Octopus maya) with baited lines in the municipality
of Campeche.”
Data availability The datasets analyzed in this study are available from
the corresponding author on request.
Declarations
Conflict of interest The author does not have relevant financial or non-
financial interests to disclose.
Ethics approval No ethical clearance was required as all octopuses
were taken by the commercial fishermen with proper fishing license.
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